United States

Department of

Agriculture

Forest Service

Northeastern

Research Station

General Technical

Report NE-288

The Role of Fire in Nongame

Wildlife Management and

Community Restoration:

Traditional Uses and

New Directions

Proceedings of a Special Workshop

Nashville, Tennessee

September 15, 2000
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The Role of Fire in Nongame Wildlife

Management and Community Restoration:

Traditional Uses and New Directions

Proceedings of a Special Workshop

Nashville, TN

September 15, 2000

Editors:

W. Mark Ford, USDA Forest Service, Northeastern Research Station,

Box 404, Parsons WV 26287

Kevin R. Russell, Willamette Industries Inc.,

Box 488, Dallas, OR 97338

Christopher E. Moorman, Department of Forestry,

North Carolina State University,

Box 8003, Raleigh, NC 27695

Sponsored by:

Tall Timbers Research Station, Tallahassee, FL

USDA Forest Service Northeastern Research Station, Newtown Square, PA

Conducted by:

The Wildlife Society, Bethesda, MD

Published by:

USDA Forest Service

Northeastern Research Station
Contents

Preface

W. Mark Ford, Kevin R. Russell, and Christopher E. Moorman ............................................................... 1

Fire in the Eastern United States: Influence on Wildlife Habitat

D. H. Van Lear and R. F. Harlow ........................................................................................................ 2

The Historical Foundations of Prescribed Burning for Wildlife: a Southeastern Perspective

A. Sydney Johnson and Philip E. Hale ................................................................................................ 11

Fire, Herbicide, and Chainsaw Felling Effects on Arthropods in Fire-Suppressed Longleaf

Pine Sandhills at Eglin Air Force Base, Florida

Louis Provencher, Krista E. M. Galley, Andrea R. Litt, Doria R. Gordon,

Leonard A. Brennan, George W. Tanner, and Jeffrey L. Hardesty ........................................................... 24

Responses of Amphibians to Fire Disturbance in Pacific Northwest Forests: a Review

R. Bruce Bury, Donald J. Major and David Pilliod .............................................................................. 34

The Effects of Burning and Grazing on Survival, Home Range, and Prey Dynamics of the

Texas Horned Lizard in a Thornscrub Ecosystem

Anna L. Burrow, Richard T. Kazmaier, Eric C. Hellgren, and Donald C. Ruthven, III ............................. 43

Fire and Bird Communities in the South

James G. Dickson ............................................................................................................................. 52

Burning for Birds: Concepts and Applications

R. Todd Engstrom and David J. Brownlie ............................................................................................ 58

Management Options for Songbirds Using the Oak Shelterwood-Burn Technique in Upland

Forests of the Southeastern United States

J. Drew Lanham, Patrick D. Keyser, Patrick H. Brose and David H. Van Lear ...................................... 65

Influence of Ecosystem Restoration for Red-cockaded Woodpeckers on Breeding Bird and

Small Mammal Communities

Ronald E. Masters, Christopher W. Wilson, Douglas S. Cram, George A. Bukenhofer

and Robert L. Lochmiller................................................................................................................... 73

Fire, Habitat Structure and Herpetofauna in the Southeast

Cathryn H. Greenberg ...................................................................................................................... 91

Numerical Response of Small Vertebrates to Prescribed Fire in California Oak Woodland

Justin K. Vreeland and William D. Tietje ......................................................................................... 100

Fire Management and Research for Biodiversity in the Green Swamp

Margit A. Bucher and Maura E. High ............................................................................................... 111

The Role of Non-industrial Private Forest Lands in the Conservation of Southern

Fire-dependent Wildlife

Christopher E. Moorman, Peter T. Bromley, Mark A. Megalos, and David Drake .................................. 116

Herbicides as an Alternative to Prescribed Burning for Achieving Wildlife

Management Objectives

T. Bently Wigley, Karl V. Miller, David S. deCalesta, and Mark W. Thomas ......................................... 124

Fire and Bats in the Southeast and Mid-Atlantic: More Questions Than Answers?

Timothy C. Carter, W. Mark Ford and Michael A. Menzel ................................................................. 139
Wildlife, Habitat, and Prescribed Fire in the Southeastern Coastal Plain: Past, Present,

and Future

Leonard A. Brennan ....................................................................................................................... 144

The Role of Fire in Riparian Zones of the Northern Rocky Mountains

Elaine K. Sutherland and Kevin McKelvey ........................................................................................ 144

The Restoration of Oak-Hickory Forests in the Central Hardwoods: Results of a

Landscape-scale Prescribed Burning Experiment in Ohio

Elaine K. Sutherland and Todd F. Hutchinson ................................................................................... 145

Fire Ecology of Marshes and Canebrakes in the Southeastern United States

Cecil Frost ..................................................................................................................................... 145
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 1

Preface

W. Mark Ford, Kevin R. Russell, and Christopher E. Moorman

Fire has a long history of regional use in the United States for forest, range and

game management. Except for a few high-profile threatened, endangered, and

sensitive species such as the pine barrens treefrog (Hyla andersonii), the redcockaded woodpecker (Picoides borealis), and the Kirtland’s warbler (Dendroica

kirtlandii), the use of prescribed burning for nongame management efforts has

been rare. Moreover, in areas such as the central hardwood regions, the

Appalachians and the Pacific Northwest, silvicultural uses of fire have been

discouraged, which consequently have limited applications of fire for nongame

management. However, new emphases on biodiversity and community

restoration are leading to re-examination of fire as a management tool in regions

with fire-dependent systems, such as the southeastern Coastal Plain and in areas

where fire historically has been excluded.

The timeliness and need of this symposium at the seventh annual meeting of

The Wildlife Society was very apparent. As forest and land management on

public lands continues to move from utilitarian or multiple-use approaches to

more ecologically holistic management programs, prescribed burning is

becoming an increasingly acceptable land management tool even where rarely

used before. Conversely, on private lands, smoke management and liability

issues and increased ownership fragmentation are having the opposite effect—

decreasing the use of prescribed fire in land management activities. As natural

resource scientists and land managers, not only must we gain a better

understanding of the effects of natural and prescribed fire (or lack thereof) on

nongame species, but also we need to promote its use when clearly indicated as

a prescription for natural resource management. A critical component of this

promotion is the education of the public by forestry and wildlife professionals

about the historical role of fire and the importance of prescribed burning as a

land management tool.

This symposium attempted to synthesize the widely scattered information and

disparate databases that exist regarding the role of fire for managing nongame

wildlife and relictual communities. Accordingly, this symposium drew upon the

expertise of regional and nationally recognized wildlife scientists, resource

managers, and fire ecologists with papers presented on the historical

background, current issues, ongoing management efforts, and research and

management needs of prescribed burning for both eastern and western

landscapes.
2 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Abstract.—Fire is a major influence shaping wildlife

habitats in the eastern United States. Lightning- and

Indian-ignited fires burned frequently and extensively

over the pre-Columbian landscape and shaped the

character of numerous ecosystems. Depending upon the

frequency, intensity, and severity of the fires, various

assemblages of plants developed along environmental

gradients, creating a shifting mosaic of habitats for

wildlife. For millennia, fire was a major ecological

process, mostly burning as frequent, light to moderate

intensity surface fires in some ecosystems and as intense,

stand-replacing fires in others. Within the past 100 years,

fire has been excluded from most of the East and firemaintained habitats have dramatically declined. In

many cases, wildlife species dependent on these habitats

are in decline or experiencing dangerously low numbers.

Unfortunately, the trend toward a relaxation of bans on

burning may soon be reversed due to the adverse

publicity from recent disastrous fire in the West.

Wildfires and escaped prescribed fires frequently have

been in the news in recent years, e.g., those in

Yellowstone National Park in 1988, Florida in 1998, and

throughout the West in 2000. Usually, only catastrophic

wildfires or escaped prescribed fires are deemed

newsworthy. Media coverage often concludes that forests

have been destroyed by these wildfires and imply that

wildlife habitat has been destroyed as well. But has

wildlife habitat really been destroyed? Does fire so

damage ecosystems that they are gone forever? Of course

not, especially in the eastern United States where

vegetative recovery is relatively rapid following

disturbance. From an ecological perspective, ecosystems

and their associated wildlife habitats are not destroyed,

only changed to an earlier stage of succession.

Fire should be considered in the context of its role as a

natural ecological process that has shaped vegetative

patterns in most eastern ecosystems for millennia. Even

ecosystems burned with high intensity fires generally

have mechanisms in place to recapture solar energy and

recycle nutrients, allowing rapid ecosystem renewal. All

over the world, early man used fire for his benefit.

Properly used, it remains an important tool in both

forest and wildlife management.

The earth has been described as a fire planet, a planet

whose terrain is covered with combustible organic fuels

which continually accumulate over time, an oxygenated

atmosphere that supports combustion, and two potent

ignition sources–lightning and man (Pyne et al. 1996).

Most ecosystems on earth have burned in the past and

will burn again in the future–creating, changing, or

maintaining habitats for a wide variety of wildlife.

Because fire was historically so prevalent and frequent

across the eastern landscape, wildlife evolved

adaptations to fire in order to survive (Komarek 1974,

Brennan et al. 1998).

Fire History in the Eastern United States

Approximately 12,000 years ago, man first made his way

into North America (Williams 1989, Bonnicksen 2000).

Within a relatively short period of less than 1000 years,

these early Americans had occupied nearly all of the

western hemisphere. They used fire in many ways

because it helped them survive. Over thousands of years,

the American Indian became expert in using fire for

various purposes, e.g., for hunting, to concentrate prey

species in convenient areas, to encourage fruit and berry

production, to keep the woods open along major

corridors of travel, to fire-proof their villages, and other

uses (Williams 1989, Pyne et al. 1996, Bonnicksen

2000). Through trial and error over millennia, the

Indian learned to make fire his servant and not his

master. Thus, controlled, or prescribed, burning is

nothing new (Komarek 1974).

The Indian population of North America at the time of

Columbus was estimated by Dobyns (1983) at about 18

million. This figure has been widely quoted, although

Stannard (1992) more recently suggested that there may

have been as many as 100 million Indians in North

America. Regardless of the accuracy of these and other

estimates of the Indian population, there can be no doubt

that, at the time of Columbus, there were millions of

Indians modifying the “natural” forests. Their influence

on the eastern forest was widespread because of their use

of fire and extensive land clearing for agriculture (Hudson

1976, Williams 1989). Because of their farming and

burning activities, Indians ensured that much of the

eastern forests was in early successional habitats.

Indians often burned as frequently as twice a year,

complementing lightning as an ignition source. Their

burning extended the fire season beyond the “natural”

lightning-fire season of summer. These frequent and

often extensive fires, along with the wildlife foraging

that fire encouraged, created and maintained open

woodlands, savannahs, and prairies throughout the

eastern United States (Williams 1989, McCleery 1993,

Kay 2000). In fact, much of the eastern forests at the

time of Columbus could be regarded as a cultural

artifact of Indian activities.

Fire in the Eastern United States: Influence on Wildlife Habitat

D. H. Van Lear1 and R. F. Harlow2

1

Department of Forest Resources, Clemson University,

Clemson, SC 29634

2Deceased; Southern Research Station, USDA Forest Service,

Clemson, SC 29634
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 3

Because of burning and agriculture, the eastern United

States at the time of Columbus was no “forest primeval”

composed of closed canopy, old-growth forests. The oftrepeated phrase that a squirrel (Sciurus sp.) could travel

from the Atlantic coast to the Mississippi River in preColumbian times without ever touching the ground is

patently false (Buckner 1983, Hamel and Buckner

1998). The eastern forest at that time was a shifting

mosaic of woodlands, savannahs, forests, and prairies,

all in varying stages of succession. After Indian

populations plummeted in the 16th century from

exposure to European diseases, their level of burning

declined and prairies gradually became woodlands,

savannahs became dense forests, and dense

undergrowth invaded open forests (McCleery 1993).

The early European settlers found that much of the East

was already being fired on a frequent basis by Indians

(Komarek 1974). These settlers gradually displaced the

Indians, but continued their use of fire for many of the

same reasons, i.e., to clear the woods of underbrush, to

expose nuts, to clear agricultural fields, etc. Frequent

fires occurred over large areas of the eastern landscape

into the early decades of the 20th century.

Not until the early 1900s were there serious efforts to

exclude fire as an ecological process in eastern North

America. In 1910, an outbreak of wildfires in the western

United States--concentrated in Idaho and Montana,

burned millions of acres, much of which was on

national forests, and killed 78 firefighters (Pyne et al.

1996). These events occurred just after the Forest Service

had made fire control a top priority on the nation’s

national forests and caused the public to become

concerned about the sustainability of the nation’s

natural resources. Led by the Forest Service and the state

forestry commissions, the public began to see fire as an

enemy to be suppressed at all costs. An era of fire

suppression began that created different environments

from those that had existed for millennia, often to the

detriment of many wildlife species.

Habitats and Fire

Habitats are environments that provide the necessities of

life for wildlife, i.e., food, water, cover, and places to rear

their young, and are shaped by ecological processes

acting on varying gradients of soil, topography,

hydrology, and climate over time. In most of the eastern

United States and in many other parts of the world, fire

historically played a major role in shaping habitats. Fire,

often working in combination with other disturbance

factors, was the major factor determining the

composition, structure, and pattern of vegetation that

occupied different portions of the landscape.

Fire became a more important force in shaping habitats

in the East as the climate warmed following the last

glaciation because a warmer climate promoted better

burning conditions. Indian populations were also

increasing, especially in the past few thousand years, as

greater use of agriculture allowed them to discard their

previous nomadic lifestyle. And wherever there were

Indians, there was fire.

The Laurentide ice sheet, which covered the northeastern

United States and extended south to the Ohio River at

the peak of the Wisconsin glaciation, began to retreat

about 18,000 years ago (Bonnicksen 2000). At the end

of this ice age about 10,000 years ago, vegetation in the

eastern United States was still migrating northward

(Delcourt and Delcourt 1987). It is difficult, if not

impossible, to unravel effects of a changing climate

(generally warming until about 6,000 years ago, then

generally cooling again–until about 1970) from the

effects of Indian burning. By about 5-6,000 years ago,

the vegetative associations of today were essentially in

place in the South, although the structure of those

forests must have been much different than today’s

forests. In the Northeast and Midwest, it took longer for

individual species to arrive in their current locations

because they had farther to travel from their southern

refugia.

Burning altered the structure, composition, and pattern

of vegetation. Fire regimes created types of vegetation

that actually encouraged recurring fire, e.g., the longleaf

pine (Pinus palustris)-wiregrass (Aristida) ecosystem, and,

in so doing, shaped habitats. The habitats that

developed depended upon the type of fire regime, the

adaptations of plants occupying the landscape, and the

length of time since the last disturbance. The magnitude

of the historical role of fire is under-appreciated by the

general public and even some resource managers

because of almost a century of efforts to exclude it from

the majority of the landscape.

Historical Fire Regimes

In the eastern United States, some areas were burned so

infrequently, e.g., moist to wet areas in northern

hardwood forests, that it can be said that these areas do

not burn. Other areas were subject to annual fires that

burned with low intensity, i.e., low flame lengths, and

low severity, i.e, little damage to soil or overstory

vegetation. Still other areas burned infrequently but

with great intensity and killed essentially all overstory

trees. Because of the many types of fires that can occur,

fire scientists have devised systems for characterizing

them (Table 1), using variables such as fire intensity,

severity, and frequency of burning to describe fire

regimes (e.g., Heinselman 1981).

Understory fire regimes are characterized by frequent

burns of relatively low intensity that do little damage to

soil or overstory trees (low severity). Ecosystems

subjected to frequent understory burns recover quickly

as the grasses, forbs, and understory woody vegetation

characteristic of this type of fire regime resprout

vigorously within weeks after burning. Overstory trees
4 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

are generally little affected by these low intensity fires.

Such a fire regime was typical of the longleaf pinewiregrass ecosystem that dominated up to 90 million

acres of the Coastal Plain of the Southeast at the time of

Columbus, but which now occupies only about 3% of

its original range (Frost 1998, Landers et al. 1995). As a

result of loss of this open woodland, savannah type

habitat, numerous wildlife species including the redcockaded woodpecker (Picoides borealis), gopher tortoise

(Gopherus polyphmus), and southern fox squirrel (Sciurus

niger) are in decline.

Stand-replacement fire regimes are typified by

infrequent fires of high intensity that kill most of the

overstory trees and sometimes damage the soil (high

severity). Such fire regimes are typical of sand pine

(Pinus clausa) stands in Florida (Outcalt and Balmer

1983), Table Mountain (Pinus pungens)/pitch pine (Pinus

rigida) stands in the southern Appalachians (Clinton et

al. 1993), and jack pine (Pinus banksiana) stands in the

boreal forests of the northern United States and Canada

(Rudolph 1983, Bonnicksen 2000). Fires in these types

may burn as surface fires under mild conditions but

under severe weather and fuel conditions standreplacement fires develop. In the absence of fire,

especially on better sites, the species are replaced by

more shade-tolerant species, i.e., scrub oaks in sand pine

(Outcalt and Balmer 1983), oaks and mountain-laurel

(Kalmia latifolia) in Table Mountain/pitch pine stands

(Waldrop and Brose 1999), and red pine (Pinus

resinosa), white pine (Pinus strobus), and various

hardwoods in jack pine stands (Rudolph 1983).

Following stand-replacement type fires, seeds from

serotinous cones germinate on a fire-prepared seedbed

and a new stand develops. However, ecosystem recovery

may take decades before the character of the new stand

recovers to that of the pre-burned stand. Burning in

these types requires intensive, experienced supervision

because of the high risk of fires getting out of control.

The jack pine ecosystem is one of the most extensive in

North America and exists in a variety of successional

stages on many site types. Jack pine grows best on dry

soils where fires are relatively frequent, i.e., every 50 -

100 years, and severe (Bonnicksen 2000). There are

many wildlife species that are characteristic of these

habitats, although we know of few that are threatened or

endangered. Many species are cyclic in the jack pine

ecosystem, including the snowshoe hare (Lepus

americanus), Canada lynx (Lynx canadensis), and the

snowy owl (Nyctea scandiaca). The Kirtland’s warbler

(Dendroica kirtlandii), one of America’s rarest birds, nests

only in pure stands of jack pine (Hunter 1990).

The Table Mountain/pitch pine ecosystem in the

southern Appalachians is in decline because of fire

exclusion in this type. There is some debate among fire

scientists about whether the Table Mountain/pitch pine

type in the southern Appalachians was maintained

primarily by stand-replacement fires or periodic

understory fires. Recent research by the North Carolina

Wildlife Resources Commission, the U.S. Forest Service,

and Clemson University suggests that periodic burning

in the type creates open woodland conditions, which

would enhance landscape diversity within the sea of

mature, dense hardwood and pine forests that now

dominate the southern Appalachian mountains.

Not all stand-replacement fires occur in forests. The

tallgrass prairie was dominated by fire-tolerant tall

grasses, including big bluestem (Andropogon sp.), little

bluestem (Schizachyrium scoparium), and Indiangrass

(Sorghastrum nutans), but also contained forbs, shrubs,

and trees. This ecosystem, which existed in pure form in

Table 1.—Heinselman’s (1981) classification of historic fire regimes, slightly modified, with selected

eastern examples. (Note: Only those classes applicable to eastern conditions are given.)

Class Characteristics of Fire Regime Eastern Examples

0 No natural fires Wetter regions of the eastern deciduous forest

1 Infrequent, light surface Many eastern deciduous forests

fires with more than 25-yr

return intervals

2 Frequent, light to moderate surface Southern pine forests of Coastal Plain

fires with 1- to 25-yr return interval and lower Piedmont; Tall-grass prairies

3 Infrequent, severe surface fires with Dry jack pine-oak “barrens” of

more than a 25-yr return interval Minnesota, Wisconsin, and Michigan

4 Short return interval crown fires and Most boreal forests in eastern Canada

severe surface fires in combination and United States; Sand pine

 (25- to 100- year return intervals)
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 5

eastern Nebraska, Illinois, Iowa, and eastern Kansas,

graded into a forest-prairie complex to the east, and has

been almost totally converted to agriculture (Telfer

2000). Only about 1% of the tallgrass prairie remains. It

was a fire-maintained ecosystem that was burned

primarily by man at intervals of 1-3 years (Pyne et al.

1996). When fires became less frequent, the eastern

portions of the prairie succeeded to shrubs and trees.

When burned annually, shrub species were eliminated.

Wildlife species in decline because of this vanishing

habitat include the greater prairie chicken (Tympanuchus

cupido), badger (Taxidea taxus), and false map turtle

(Graptemys pseudogeographica).

Some fire regimes may include both surface fires and

stand-replacement fires. There are mixed severity fire

regimes where fires alternate between understory burns

and stand replacement burns, either within the same fire

or in different fires. Such burns cause selective mortality

in the overstory, especially when certain species are

better adapted than others to withstand the effects of

high intensity fires. Burns of this type occur, for

example, in mixed pine/hardwood stands in the

southern Appalachians where fuel loadings have

accumulated primarily in the form of dense understories

of mountain-laurel (Elliott et al. 1998, Waldrop and

Brose 1998). Fires in this type are intense enough to

topkill many of the hardwoods and shrub understory

but the thick bark, high crowns, and ability to quickly

regenerate new needles allows many of the mature pines

to survive.

Fire Adaptations

Fire is only one of many evolutionary forces that shape

plants and plant communities. However, fire has been

so ubiquitous on the planet and exerts such profound

influences on the environment that it is reasonable to

assume that fire regimes selected for individuals in

populations that could best survive fire (Pyne et al.

1996). Plant species that evolved with fire adapted by

developing characteristics that increase their chances of

survival in various types of fire regimes (Table 2). Such

adaptations occur in various stages of a plant’s

development, from regeneration to maturity (Kimmins

1987).

The ability to resprout from root collar buds following

topkill by fire is probably a fire adaptation in some

species, even though this characteristic occurs in other

taxa where fire is alien (Bond and van Wilgen 1996).

Many species replace scorched foliage by sprouting from

adventitious or latent axillary buds on branches or boles

of trees. Where fire is frequent, sprouting adaptations

Table 2.—Plant adaptations to fire in different stages of development.

Stage of Development Adaptation Plant Examples

Vegetative Sprouting from root collar buds Most hardwoods;

certain pines

Sprouting from adventitious or latent Most southern pines

axillary buds

Thick bark Most oaks; southern yellow pines;

numerous others

Protected buds on boles and limbs Pitch pine and pond pine

Protected buds below ground Oaks

Reproductive Precocious flowering Table Mountain pine;

Sand pine; jack pine

Stimulation of flowering Wire grass

Serotinous cones Table Mountain pine; pitch pine;

jack pine; sand pine

Germination Heat-enhanced seed germination Some legumes; fire cherry;

yellow-poplar

Germination on fire-prepared seedbeds Many pines; fireweed;

many other species

Other Adaptations Increased inflammability Longleaf pine, wiregrass, jack pine
6 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

such as these improve the survival potential of plants

because it allows those plants to continually occupy the

same growing space. Non-sprouters would be favored by

longer return intervals between fires because they could

allocate more of their energy to top growth, allowing

them to more completely occupy the growing space in

the absence of fire for long periods.

Some species have developed thick bark apparently to

insulate the living cambium from the heat of surface

fires. Other species have thin bark when young and are

quite susceptible to fire damage or mortality, but

develop very thick bark as they mature and become

resistant to surface fires. The protection offered by bark

is dependent upon not only its thickness but, at least for

some species, by its thermal properties (Hare 1965).

Certain fire-adaptations are even more species specific.

Cone serotiny, where seed accumulates in unopened

cones that open after being heated by fire, is

characteristic of relatively few species in the East. Species

with serotinous cones normally exist in environments

where high intensity, stand-replacing fires are the norm

(Pyne et al. 1996, Bond and van Wilgen 1996). Another

species-specific adaptation is the grass stage of longleaf

pine, where their fire-sensitive buds are protected from

overheating by a dense sheath of moisture-rich needles

(Walhenberg 1964). This characteristic is unique among

the pines.

Oaks in some ways might be considered to be the

longleaf pine of the hardwoods. They have multiple

adaptations to fire, including thick bark that protects

them from damage by surface fires. Oaks also have

hypogeal germination, i.e., their root collar buds are

below the surface of the soil because their acorns are

buried by squirrels and/or jays (Sciuridae and Corvidae).

Because these buds are protected from the heat of

surface fires (soil is a poor conductor of heat), oaks are

better able to successfully sprout following fire than

many of their competitors whose seeds germinate on the

soil surface, such as yellow-poplar (Liriodendron

tulipifera) (Brose and Van Lear 1998, Brose et al. 1999).

Many species in the eastern United States have seeds that

are either encouraged to germinate by heating or to

germinate on fire-prepared seedbeds (Pyne et al. 1996,

Lyon et al. 2000). Heat from fire can crack the seedcoat

and release the seed from dormancy. Some grasses, such

as the wiregrasses, only flower when burned in certain

seasons, with blooming following spring burns but not

after winter fires (Brennan et al. 1998).

Some adaptations not only allow plants to survive fire

but also play a major role in predisposing plant

communities to recurrent fire (Mutch 1970). For

example, the dominance of wiregrass in the herbaceous

layer, with its high proportion of dead leaves and their

overlapping nature, along with the resinous nature of

fallen longleaf pine needles, insures that surface fires

will be frequent in this ecosystem (Landers et al. 1995).

Another example would be in oak forests where the

fallen leaves curl and resist decay, allowing a highly

aerated fuel bed to develop, which encourages frequent

surface fires. In contrast, leaves of fire-sensitive

hardwoods like American beech (Fagus grandifolia),

yellow-poplar, and red maple (Acer rubrum) lie flat on

the ground and decay rapidly, preventing a flammable

fuel bed from developing. Unless fire-adapted plant

communities tend to promote recurrent fire, they are

likely to be replaced by non-flammable communities

(Bond and van Wilgen 1996).

These adaptations of different plant species, along with

conditions for establishment and the availability of

plant propagules, determine patterns of succession and

the types of habitats that develop following burning.

Effects of Fire on Habitat Components

Fire affects habitat components in many ways. Periodic

understory burns cause many plants to resprout and

these new sprouts, whether woody or herbaceous, are

more palatable and have higher protein and nutritive

content than the older tissues that they replaced

(Stransky and Harlow 1981, MacCracken and Viereck

1990). Enhanced palatability and higher nutrient

contents in post-burn growth are relatively short lived

(generally 1-2 years), but are important in areas

frequently burned. Regrowth after burning is not only

more palatable and nutritious than older tissue, it is also

more available because most sprouting occurs at ground

level (Lay 1967).

Frequent burning tends to favor herbaceous vegetation

over woody vegetation, i.e., it increases the proportion

of herbage to browse (Lewis and Harshbarger 1976,

Pack et al. 1988). Growing season burns discriminate

against woody growth more than dormant season burns.

Frequent burning creates open forest conditions in both

pine and hardwood forests, i.e., understory and

midstory woody vegetation are replaced by herbaceous

vegetation (Landers et al. 1995, Thor and Nichols

1974). Burning does not eliminate woody growth but

controls its size (Waldrop et al. 1987). Annual summer

burns in southern pine stands will eventually remove

hardwoods from the stand but it takes numerous burns.

Certain woody species, such as oaks, tend to be more

tenacious sprouters than competitors and therefore

persist in frequently burned ecosystems.

Seed and mast production generally increase after

burning in Southern forests. Seed production by

legumes, grasses and spurges is usually greater on

frequently burned areas (Cushwa et al. 1970, Robbins

and Myers 1992). Production of berries, drupes, and

pomes normally peaks about 2 to 4 years following

burning (Harlow and Van Lear 1989). However, burning

too frequently, i.e., annually or biennially, reduces most

mast-producing species (Robbins and Myers 1992).
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 7

Numerous hard-seeded species require their seed to be

heated to 60 - 80o

C to break dormancy (Auld 1986).

Some shrub species (Vaccinium) and small trees such as

dogwood (Cornus florida) tend to be more fruitful a few

years following burning, although others do not

(Harlow and Van Lear 1989). Hard mast-producing

species, such as oaks and hickories (Carya sp.), are

generally favored by fire during the regeneration period

(Abrams 1992, Barnes and Van Lear 1998, Brose et al.

1999).

Effects of fire on standing dead trees, i.e., snags, and

fallen woody debris are variable (Lanham and Guynn

1993, Van Lear 1993). Surface fires burning under

droughty conditions frequently kill stressed trees,

creating snags. In addition, fires may scar the bases of

some trees and eventually become basal cavities.

However, some snags may be consumed by fire or at

least brought to the ground, where they become fallen

woody debris. Both forms of coarse woody debris are

extremely important habitat components (McMinn and

Crossley 1993). In the East, most snags do not stand for

long periods of time, often falling within a decade of

death (Dickenson et al. 1983, Sabin 1991). Coarse

woody debris is generally not consumed by fire unless

the drought index, e.g., the Keetch-Byrum drought

index, is high. Only when the debris becomes dry

throughout during droughty conditions will significant

amounts be consumed by fire.

In the southeastern United States, long-term studies in

the Coastal Plain show that periodic burning increases

availability of phosphorus, as well as other nutrients in

the mineral soil, probably because of increased

mineralization and because soil pH is raised (McKee

1992). Most soil nutrients become more available for

plant uptake as soil acidity approaches neutral. This

increased availability probably accounts, at least in part,

for the higher nutrient levels in regrowth following fires.

Enhanced soil phosphorus levels resulting from burning

may have implications in Quality Deer Management

programs because phosphorus is needed for antler

development (Cowan and Long 1962).

Burning historically provided a mosaic of habitat types

in a variety of seral stages and fuel conditions over the

landscape (Wood 1981, Williams 1989, Pyne et al.

1996). Fire did not burn uniformly across the landscape,

but interacted with topographic, edaphic, vegetative, and

weather conditions to produce a mix of habitats and

ecotones. While fire occasionally changed mature forests

to early successional habitats, i.e., jack pine ecosystems,

it also maintained the structure and composition of

certain ecosystems where fire was a frequent visitor, e.g.,

the longleaf pine-wiregrass ecosystem and the tallgrass

prairie ecosystem. These varied conditions created

habitats for many species of wildlife which, over

millennia, came to depend on these conditions. As the

role of fire has declined over the past century, so have

many plant and wildlife species.

Conclusions

Fire has shaped the pattern, composition, and structure

of vegetation in the eastern United States for millennia,

especially since the arrival of Indians at least 12,000

years ago. Indians used fire for many purposes and often

fired the woods on an annual basis. Frequent burning

kept the forests more open and increased the ratio of

herbaceous/woody vegetation. Fire did not burn

uniformly across the landscape but created a shifting

mosaic of habitats depending upon fuel, weather, and

site conditions at the time of burning. In some

ecosystems, fire burned frequently but with low

intensity and severity and maintained the status quo of

the ecosystem. In other cases, fires burned with high

intensity and severity and completely changed the

ecosystem. Plants and animals adapted to these different

fire regimes in order to survive. Over time, many wildlife

species became dependent upon these fire-maintained

habitats.

During most of the past century, attempts have been

made to exclude fire from much of the eastern

landscape. Exclusion of fire, along with other factors,

have resulted in major declines in fire-maintained

habitats, the most notable examples being the longleaf

pine-wiregrass habitat and the tallgrass prairie habitat.

Fire also played a role in maintaining other habitats

important to wildlife. Historically, it perpetuated the

jack pine, sand pine, and Table Mountain/pitch pine

ecosystems and was instrumental in maintaining oak

forests on good-quality sites throughout much of the

East. The exclusion of fire in fire-maintained habitat

types may make catastrophic wildfires more frequent,

potentially changing ecosystems and eliminating

desirable habitat for some species. Burning on a regular

basis prevents fuel build-up and minimizes the chances

of a catastrophic wildfire occurring.

There is no question of the historical importance of fire

in creating and maintaining wildlife habitats in the

eastern United States. However, we now live in a

different time and there are numerous restrictions on

the application of fire for wildlife habitat purposes. We

suggest that prescribed fire be used where feasible and

where it can accomplish desired objectives, because it

has unique habitat effects. However, in the future it will

be necessary to use other management techniques,

including harvesting, thinning, and herbicides, to

mimic, at least partially, the effects of prescribed fire on

wildlife habitat where burning is not appropriate.

Acknowledgments

This manuscript benefitted from reviews by Robert

Downing and Bently Wigley.
8 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

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The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 11

Abstract.—Controlled burning has deep historical roots

in the South, where the practice was quickly adopted

from the Indians by early European settlers. It became

used widely, primarily to improve forage conditions for

free-ranging cattle and to improve visibility and access.

Likewise, hunting is deeply imbedded in southern

culture and was an attraction to visitors throughout the

19th Century. This was especially true of quail (Colinus

virginianus) hunting, and after the Civil War wealthy

northerners began to buy large plantations for hunting

retreats. In the 1920’s Herbert L. Stoddard documented

the necessity of prescribed burning to maintain

bobwhite quail habitat on these plantations. Opposition

to the practice among foresters and public agencies was

fierce, and Stoddard became an outspoken advocate of

light winter burning in longleaf pine (Pinus palustris)

and, later, certain other forest types. Use of prescribed

fire in forestry and game management was gradually

accepted. But, although some naturalists such as

Stoddard were interested in the effects of fire on native

flora and nongame wildlife, private landowners and

public agencies generally showed little interest in

managing specifically for non-game wildlife until the

1970’s. By then, there was in the southern states a

background of 50 years of research and many more years

of practical experience in the use of fire that could be

applied to this new goal. Soon, any biologists and

managers recognized that prescribed burning would

play a nearly essential role in managing certain

nongame species. And, as new management goals

evolved, fire regimes other than light winter burning

also came under scrutiny for potential use in restoration

and maintenance of certain natural communities

Introduction

Woods burning has a long unbroken history in parts of

the South, and much of the early scientific work on

prescribed burning was done there. In the keynote

address to attendees of a prescribed burning

symposium, E. M. Bacon (1971) of the U.S. Forest

Service said “Prescribed burning seemingly had its origin

in the South, it has certainly been an accepted

management tool for a longer period of time in this

region than anywhere else in the country.” In his book

“Fire in America,” Pyne (1982) noted that when the

need for prescribed burning became evident, the skills

were in the hands of local woods burners, “a resource

that had to be managed.” He concluded that “Its

peculiar fire heritage helped the South to train the rest

of the nation in the art of prescribed burning.”

There is too much literature on the subject to review in

detail. The history of the controversy surrounding

controlled burning and the forestry profession has been

dealt with in detail by others. There are at least two

important books dealing with the subject (Pyne 1982;

Schiff 1962). Our objective is to provide a historical

overview of the southern roots of prescribed burning in

forestry and, especially, wildlife management. We will

mostly limit our discussion to the application of

prescribed burning in management and will not review

research on habitat and wildlife responses to fire. The

term prescribed burning originated among foresters and

came into use in the 1940’s. The term controlled

burning was used by the local people and, until recently,

by wildlife managers. Although the two terms have

somewhat different definitions, we will use them

interchangeably, as appropriate in the context of the

discussion.

Prehistoric Fires

Fire is more common and more important in the

environment of the South than in most other areas of

the United States (Christensen 1978). This is especially

true in the Coastal Plain, but even in the Appalachians

fire has been frequent enough to cause the development

of fire-dependent species and influence the composition

of forests over large areas (Van Lear and Waldrop 1989).

The frequency, seasonality and intensity of its

occurrence are determined in part by local climate,

topography, soils, and vegetation. Plants adapted to

specific sets of conditions are sorted into communities

that may have characteristics that facilitate or deter fire,

or influence the type of fire regime. Periodic fire is

especially an important part of the environment of all

the southern (yellow) pines (Pinus spp.) except spruce

pine (P. glabra), which typically grows in mixed stands

with hardwoods. Landers (1991) stated “the pine genus

has long been recognized as having special traits

adaptive to fire and fire-prone sites,” and he provided a

detailed analysis of these traits and their relationship to

fire periodicity and intensity for each of nine southern

pine species. Today, nearly half of the total forest area in

the southern states and nearly two-thirds of that in the

Coastal Plain and Piedmont is occupied by pine or pinehardwood types, (estimated from data in Rudis 1998).

But pine forests are not the only fire-adapted

communities. Other community types that are

dependent upon periodic fires for their regeneration or

The Historical Foundations of Prescribed Burning for Wildlife:

a Southeastern Perspective

A. Sydney Johnson1 and Philip E. Hale1

1

Daniel B. Warnell School of Forest Resources, University of

Georgia, Athens, GA 30602
12 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

maintenance include shrub bogs, Atlantic white-cedar

(Chamaecyparis thyoides) swamp forests, bay forests,

cypress (Taxodium ascendens) domes, and grass-sedge

bogs in the Coastal Plain (Christensen 1978, 1988;

Garren 1943; Komarek 1974,), heath thickets and grassy

meadows at high elevations in the Southern

Appalachians (Christensen 1978; Garren 1943), and

canebrakes along stream bottoms in all provinces

(Hughes 1966; Komarek 1974; Platt and Brantley 1997).

Lightning is assumed to have been the primary agent

that caused fires in North America before humans

became a major factor (Komarek 1964, 1966; Pyne

1982). Until recent centuries there were few natural

barriers to the spread of such fires after ignition, and

fires probably swept unimpeded over vast areas.

However, some (e.g., Stewart 1956) have questioned

whether lightning played the primary role in shaping

pine forests in much of the South. They have noted that

lightning in the South is usually accompanied by rain,

and lightning-caused fires in the region today are usually

spotty and small in size compared to man-caused fires

(Barden and Woods 1976).

Anthropologists and geographers tend to emphasize

man as the main source of prehistoric fire (Denevan

1992; Stewart 1956). Paleoecological studies (Delcourt

and Delcourt 1985; Delcourt et al. 1993; Whitehead and

Sheehan 1985) indicate there was a rapid expansion of

southern pine forests 3,000-6,000 years ago. This

occurred concurrently with the expansion of American

Indian populations and may have been at least partly

due to their actions. But, it is impossible to sort out the

effects of climatic changes that were occurring during

that time. Many writers have documented deliberate use

of broadcast burning by the Indians (DeVivo 1990;

Hammett 1992; Maxwell 1910; Robbins and Myers

1992; Rostlund 1957; Silver 1990; Stewart 1956; Van

Lear and Waldrop 1989). Spanish explorers and, later,

other travelers consistently described an open landscape

and gave eyewitness accounts of Indians using fire in

clearing land for agriculture, driving game, and other

purposes. Rostlund (1957) reviewed many of the early

narratives and concluded that Indians were burning the

landscape and creating savannas and prairies at an

accelerating rate until European contact in the 16th

Century. Increasing amounts of charcoal in deposits laid

down over the last 1,000 years (Delcourt and Delcourt

1985, 1997) support Rostlund’s conclusion.

After contact with Europeans, Indian populations were

severely decimated by disease and other factors—

possibly being reduced by as much as 90 percent (Lovell

1992). The prairies and savannas returned to forest, and

the extensive pine forests described by so many travelers

in the 18th and 19th centuries (Johnson 1987)

probably developed after the influence of the Indians

was greatly diminished, or removed entirely. Regardless

of the relative importance of man versus lightning as a

pyrogenic agent shaping vegetation, it is evident that for

several thousand years, fires set by Indians were

significantly affecting the landscape (Delcourt and

Delcourt 1997).

The Tradition of Woods Burning

in the South

As European settlers moved in, fire was used to clear

land for farms and communities; and, nearly everywhere

on the frontier, fire was used carelessly at first. But,

controlled burning did not become a common practice

in the Northeast. Much of the region was mesic

hardwood forest (Kingsley 1985), not a fire type, and

there were fire control laws in all of the Northeast by the

time of the Revolution (Pyne 1982). Commercial

logging in the region began early, and timber interests

encouraged fire suppression (Pyne 1982). Also, the

Northeast was settled mainly by people from the

southeastern lowlands of England (Fischer 1989), and,

later, from parts of continental Europe where forests

were predominantly fire-sensitive hardwoods or spruce

(Picea spp.). Immigrants from these areas had little

experience with controlled burning (Pyne 1982). They

came from some of the most heavily urbanized areas of

the time; few of them had lived on farms (Fischer

1989), and most that had lived on farms came from

areas where row crops and fenced pastures were the

dominant pattern of agriculture (Fischer 1989;

McWhiney 1988; Pyne 1982). They developed

agricultural patterns in America similar to those with

which they were familiar (Fischer 1989).

But, in the South, woods burning was a widespread

practice from the outset, especially in the Coastal Plain.

As previously shown, much of the region supported fire

type forests. Settlement was primarily by immigrants

from the uplands of rural western England, and later

from Scotland and Ireland (McWhiney 1988, Fischer

1989), where open range herding was customary

(McDonald and McWhiney 1975; McWhiney 1988;

McWhiney and McDonald 1985). Many of the settlers

probably were already familiar with the practice of

controlled burning to improve grazing conditions. The

forests in the highlands of England, Scotland and

Ireland had been converted to heathlands by centuries

of burning for grazing by cattle and, later, sheep and red

grouse (Gimingham 1970; Kayll 1966). In much of the

South people called “crackers” (sometimes

disparagingly) practiced a frontier lifestyle centering

around free-ranging livestock (cattle and hogs), patch

farming, dipping turpentine, and hunting and fishing

(McWhiney 1988; Owsley 1949). Pyne (1982) noted

that “for many of the Scotch-Irish immigrants who

settled the regions, the socio-economic environment

was not unlike that of Scotland, which had helped to

perpetuate a herding and hunting economy that

routinely used broadcast fire.” McWhiney and

McDonald (1985) observed “The custom of range

burning. . .seems to have been another adaptation in
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 13

America of a Celtic tradition—one that meshed with an

American Indian practice.” McWhiney (1988) stated

that open range herding of livestock was “a continuation

in the Old South of traditions practiced for centuries by

Celtic peoples.” In Florida and the Southwest, there was

a similar Spanish influence.

As settlement continued, the better lands, especially in

the Upper Coastal Plain, Piedmont and Valley provinces,

were increasingly occupied by descendants of English

Cavaliers who had moved inland from coastal Virginia

and the Carolinas. They were mainly farmers and

planters, who cleared the forests, cultivated the land and

developed permanent lifestyles (Owsley 1949). Here,

the plantation culture was best developed and, because

most of the land was cleared for row crops and

improved pasture, in many places the tradition of

burning the woods was broken. Those who were feeling

crowded and unwilling to change their ways and

become farmers, moved to less productive lands

(Owsley 1949) or on to the West (Jordan 1981). They

were able to pursue their traditional ways in the

extensive pinewoods of the Coastal Plain, the Southern

Appalachians, and scattered areas throughout (Owsley

1949). In these areas woodlands were treated as a

commons for hunting and grazing.

Popular interest in the Old South has focused mainly on

the large cotton and rice plantations, while, until

recently, the other livestock producers have gone

unrecognized except by professional historians. Unlike

the plantation owners, small farmers and others who

lived off the land left little in the way of written records.

McWhiney (1988) cited data showing that for 15 years

before the Civil War the average number of livestock

driven to market from the South each year was many

times the annual numbers in the famous Texas cattle

drives during their heyday for 15 years after the war. Just

before the Civil War, the cash value of southern

livestock equaled the cash value of cotton and all other

crops combined (McWhiney 1988). Much (but not all)

of the livestock was produced on open range. Fencing of

one’s property was illegal in some southern states until

the middle 1800’s (McDonald and McWhiney 1975);

and, in most states of the Deep South, unrestricted

ranging of livestock was allowed under local option laws

until after World War II. It was up to the landowner to

fence livestock out, and motorists were liable for

damages in accidents involving livestock on roads and

highways.

Many of the herdsmen tended large herds of cattle and

hogs but owned little or no land. Some were cowboys

hired by absentee livestock owners. In the pinewoods

large herds were tended from horseback, and controlled

burning opened up the landscape, allowing better access

and visibility. Other reasons for burning included

reducing the hazard of wild fire to turpentine woods,

reducing the risk of rattlesnake (Crotalus spp.) bite, and

controlling ticks. Often no attempt was made to confine

fires within property boundaries, nor was there a desire

on the part of most landowners to exclude fire from

their property. Controlled burning was often a

community affair.

Fire also was used as a tool in hunting, however, did not

find any references documenting prescribed burning

specifically to improve game habitat. Nonetheless,

hunting was an integral part of Southern culture among

all social and economic classes (Elliott 1846; Gohdes

1967; Marks 1991; Oppel and Meisel 1987). Skilled

hunters would have known which habitat conditions

provided best hunting success and what practices

resulted in those conditions. And they almost certainly

would have used these practices to increase hunting

success—especially on the plantations, where sportsmen

from the northern states and Europe were entertained

throughout the nineteenth century.

The Quail Plantations

and Controlled Burning

Hunting in the South, especially for bobwhite quail

(Colinus virginianus) hunting, became increasingly

attractive to northern visitors after the Civil War. As land

became available during the economically difficult times

following the Reconstruction era, wealthy northerners

began to buy plantations for hunting retreats in South

Carolina, Florida, southwestern Georgia, and, later, in

other areas of the Southeast (Brueckheimer 1979;

Paisley 1968). One of these owners, Henry L. Beadel,

told of how he and other new owners brought northern

attitudes about fire with them and were appalled at the

extensive burning (Beadel 1962). They put a stop to it

on their properties, and as a woody understory

developed, quail populations declined.

But, these plantation owners were educated and

influential. In 1923 they began discussions with the U.

S. Bureau of Biological Survey (forerunner of the Fish

and Wildlife Service) that led to a cooperative study of

the bobwhite quail and the reasons for its decline.

Herbert L. Stoddard was employed by the Bureau to

head the project, which was financed by the plantation

owners. The final report on the study was published in

book form and remains a classic in wildlife literature

(Stoddard 1931). In it, Stoddard identified lack of fire as

a cause of quail decline. He became an outspoken

advocate for the use of light winter fires in game

management and forestry, especially in the longleaf pine

(P. palustris) type. After completion of the original quail

investigation., Stoddard continued to promote

prescribed burning as director of the privately funded

Cooperative Quail Study Association from 1931 to 1943

and, after that, as a forestry and wildlife consultant, and

still later, as one of the founders of Tall Timbers

Research Station near Tallahassee, Florida.
14 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

The Controlled Burning Controversy

For years a conflict had been developing between timber

interests and local livestock interests over range burning

and free-ranging livestock, especially hogs. While

economic interests of lumber companies, foresters, and

some absentee landowners focused on timber, the

economic interests of the local people were centered on

the understory grasses for cattle grazing and, in the

pinelands, on turpentine production. As early as 1850,

the state geologist of Mississippi reportedly stated “The

beautiful park-like slopes of the pine hills are being

converted into a smoking desert of pine trunks on

whose blackened soil the cattle seek more vainly every

year the few scattered sickly blades of grass whose roots

the fire has not killed” (Dunston 1913). A U. S. Forest

Service examiner (Dunston 1913) reported three-fourths

of the woodlands in Mississippi were burned over every

year and many localities were burned twice or more. He

called for a vigorous education campaign, stating “The

enormous loss caused by the boll weevil is not greater

than that from forest fires, nor is the extermination of

the cotton pest of more vital importance to the future

wealth of Mississippi than the prevention of such fires.”

Similar conditions and concerns were reported for

pinelands elsewhere. W. W. Ashe (1894) made a survey

of forest conditions in eastern North Carolina for the

North Carolina Geological Survey and reported a chief

consideration in the regrowth of longleaf pine in North

Carolina to be “entire prevention of all fires among the

pines.” In another report on forest conditions in North

Carolina, Gifford Pinchot and Ashe (1897), stated “The

first and absolute prerequisite before any attempt can be

made to improve the condition of the long leaf pine

forests is entire exclusion of cattle and hogs and

complete protection from fire.”

The hardwood areas of the mountains were similarly

burned and grazed. Regarding forest conditions in the

mountain region of North Carolina at the turn of the

century, Holmes (1911) reported unrestricted grazing

despite local stock control laws in some counties. He

reported 20 to 50 percent of the forest land in each

county was burned annually “with the false idea that

(fires) improve the range.”

Throughout much of the South, Coastal Plain and

mountains, large lumber companies began extensive

commercial harvest of timber in the late 1800’s. The

wasteful and destructive logging pattern, typical of the

time, left behind vast amounts of slash and logging

debris. Annual burning of the cutover lands continued,

intensified by the heavy fuel loads.

From the foresters’ perspective, there was reason to be

concerned. Foresters correctly perceived that

uninterrupted annual burning did not allow forests to

regenerate. However, many of their other ideas proved to

be wrong—for example, claims that fire should be

totally excluded from longleaf pine forests (Ashe 1894,

Pinchot and Ashe 1897), that it resulted in “soil

impoverishment” (Dunston 1913), that it resulted in

short-bodied, limby, knotty longleaf pine trees of low

value (Ashe 1894), and that burning actually harmed

the grazing resource rather than improving it as the

woods-burners contended (Dunston 1913; Holmes

1911).

In this political environment it is not surprising that

Stoddard’s articulate advocacy of controlled burning was

fiercely opposed, especially by foresters. The U. S. Forest

Service maintained a firm policy against burning

(chronicled by Schiff 1962). Federal funds available for

state forestry agencies under the Clarke-McNary Act of

1924 were withheld from states if they tolerated

controlled burning. The American Forestry Association

sponsored a massive propaganda campaign, the

Southern Forestry Education Project, from 1927 to

1930. Teams of men, known as the “Dixie Crusaders”

were sent into the rural South with trucks equipped with

generators, movie projectors, films, radio broadcasts,

posters, and pamphlets. According to Schiff (1962), “the

crusaders traversed 300,000 miles, disseminating 2

million pieces of literature along the way. More than

5,200 motion picture programs and lectures were

presented to 3 million people....” Damage to wildlife

was emphasized in the campaign.

Also, beginning in the 1930’s, the Forest Service

employed sociologists and psychologists to study the

woods-burners and continued to support research on

the subject for more than 40 years. In mostly

unpublished reports to the Forest Service (summarized

by Pyne 1982), the researchers concluded that

underlying reasons and motives for woods burning

included social isolation, boredom, ritualistic tradition

(“our pappies burned the woods,” Shea 1940),

frustration of a culturally and economically

disadvantaged group, alienation, and creation of jobs in

fire suppression.

The developing pulp and paper industry added its

influence to the anti-fire effort. In the 1940’s the

National Advertising Council, the U. S. Forest Service

and state forestry agencies created what has been called

the most effective advertising campaign in history: the

Smoky Bear program. And, in an appeal to the strong

religious convictions held by most rural southerners,

compilations of Bible passages deemed to be anti-fire

were printed in pamphlets and widely distributed (e.g.

U. S. Department of Agriculture 1955) and at least one

poster showed Smoky Bear praying for people to be

careful with fire.

At the time of his original study of bobwhite quail,

Stoddard was an employee of the Bureau of Biological

Survey in the U. S. Department of Agriculture, and

publications by employees had to go through review by
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 15

other agencies in the Department that might be affected.

Stoddard had great difficulty in getting the chapter on

controlled burning in his bobwhite quail book past

reviewers in the U. S. Forest Service and had to revise it

several times (Stoddard 1969).

In his memoirs, Stoddard (1969) wrote that the anti-fire

campaign was “the most intensive—and ludicrous—

educational campaign that ever insulted the intelligence

of American audiences. It was carried on by wellmeaning but utterly misinformed persons.”

Stoddard wrote:

“One of the main arguments against the custom of

burning off the longleaf forests was the alleged

disastrous effect on game and wildlife in general. . .

Motion pictures of deer with hair and hooves burned off

in disastrous crown fires (in the west or the Lake States)

were regularly shown on southeastern screens.... Dead

fish, killed by the ash from explosive mountain fires,

were shown rotting in the sun as examples of what

would happen if the Florida cattlemen did not stop

firing the flatwoods.”

Stoddard complained that, as forestry agencies gradually

withdrew their unalterable opposition to the use of fire

“in a gesture that might be termed face-saving, they

substituted the expression ‘prescribed burning’ for

‘controlled burning,’ and insisted that experts must

‘prescribe’ the practice.”

In their own defense, foresters noted that burning was

not easily confined within property boundaries, and

they contended that advocating prescribed burning

while conducting a program to persuade people to

reduce the occurrence of wildfire would send a mixed

message and weaken fire control programs. Also, they

noted that advocates of prescribed burning did not have

the responsibility of controlling destructive fires as did

foresters. (Reibold 1971).

Acceptance of Prescribed Burning

by Public Agencies

U. S. Forest Service policies toward prescribed burning

were especially important to wildlife management

during the first half of the century because the Forest

Service at that time controlled most public land

important to wildlife in the South, including most

wildlife management areas, which were under

cooperative agreements with state wildlife agencies. It

also greatly influenced state and private actions through

funding programs and cooperative work with state

forestry agencies and educational programs.

Reibold (1971), in reviewing the history of prescribed

burning from the perspective of the Forest Service,

explained that a major reason that forestry agencies were

slow to put prescribed burning into practice was

inadequate budgets and professional staff, coupled with

the need to address other priorities. In the 1930’s, when

large-scale acquisition of national forests was

undertaken with little professional staff and inadequate

budgets, priority had to be given to hundreds of

transactions involved in land acquisition, mapping,

development of work programs for Civilian

Conservation Corps camps, reforestation, and fire

control. Also, Reibold contended that proper execution

of prescribed burning was not possible until tractor

plows became generally available, which was not until

after World War II.

But Reibold (1971) acknowledged that there was much

resistance among foresters to prescribed burning. Nearly

all of the early foresters in the South were from the

North and the West. A strong German protectionist

influence in the forestry schools was greatly reinforced

by experiences with the extremely destructive wildfires

that followed logging in the northern states, burning

millions of acres and, in several cases causing hundreds

of human deaths. Reibold said “some (foresters), having

spent a lifetime in working to prevent fires or in fighting

them, were emotionally opposed to what seemed to

them to be an abandonment of all they had worked

for.”

Early in the Twentieth Century, some scientists and

professional foresters had begun to recognize the role of

fire in maintaining longleaf pine forests. The

contributions of a few individual botanists, foresters and

animal husbandrymen, who bucked the tide, are

detailed by Harper (1962), Pyne (1982) and Schiff

(1962). Although administrators reportedly suppressed

and delayed publication of research on controlled

burning (Pyne 1982; Schiff 1962), some research

personnel in the U. S. Forest Service were reporting on

the effects of burning and its possible use in silviculture.

A listing and abstracting of publications of the Southern

and Southeastern Forest Experiment Stations from 1921

to 1955 (Bruce and Nelson 1957) shows publications

dealing with the benefits of prescribed burning in forage

production, pine regeneration, fire hazard reduction,

control of brown spot needle rust, and hardwood

control appearing as early as 1932 and becoming

numerous in the 1940’s. There were proposals and

directives for limited prescribed burning on federal

lands in the 1930’s (Reibold 1971). But, with a few

exceptions, research findings and policy directives were

not implemented, and prescribed burning for the most

part was restricted to private lands. There was little use

on public lands until the 1960’s, and even the forest

industries were reluctant to do much prescribed

burning.

It was not until after World War II, when equipment and

manpower became available that the national forests

began to undertake burning, mostly for fuel reduction.

In 1947 prescribed burning was initiated on the Francis
16 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Marion National Forest, including some growing season

fires for hardwood control and periodic winter fires to

improve habitat for wild turkeys (Meleagris gallopavo)

(Devet and Hopkins 1967; Holbrook 1952). This was

one of the earliest cases of prescribed burning on a

national forest with wildlife habitat improvement as an

objective. By the decade of the 1950’s, prescribed

burning on the national forests averaged about 250,000

acres per year (Reibold 1971). During the 1960’s, the U.

S. Forest Service cautiously began expanding prescribed

burning on forests outside the longleaf pine region.

In 1971 the U. S. Forest Service sponsored a symposium

on prescribed burning. In the foreword to the

proceedings (USDA 1971) symposium chairman

Stephen Boyce stated that among the 450 scientists, land

managers, and environmentalists attending, “consensus

was essentially unanimous that prescribed fire, when

properly used in the South, is an almost indispensable

management device having generally beneficial effects,

certainly lacking in sustained deleterious effects on the

crop trees, on the soils on which they grow, or on the

flora and fauna of the area burned.” Symposium

participants, however, did recognize a lack of knowledge

about the effects of prescribed burning on air quality—a

matter of growing concern.

Prescribed burning had not been as big an issue with

other federal agencies because they controlled few areas

with upland forests in the South. There were only four

national wildlife refuges with large areas of upland

forest. There was some prescribed burning on the

refuges in the 1940’s, but generally for uplands there

was a fire exclusion policy from 1949 until even-aged

forest management was adopted in 1962 (Czuhai 1981;

Givens 1962). Prescribed burning in waterfowl marshes

(Lynch 1941) had been accepted much more readily

than on forestland (Givens 1962).

The National Park Service had a strong anti-fire policy

from the time of its establishment in 1916 until 1958

when a policy change was made specifically for the

Everglades National Park and the first prescribed burn in

a national park was conducted that year. The antifire

policy of the National Park Service was reversed in 1967

(Hendrickson 1972; Kilgore 1974; Taylor 1981) to allow

prescribed burning where necessary for community

restoration and maintenance.

At the state level, most wildlife management areas were

not owned by the states but were operated under

cooperative agreements with landowners, who approved

or disapproved habitat management measures. Before

about 1960, most management areas in the Southeast

were on national forests, many of them in the

mountains. Thus, state wildlife agencies had little

opportunity to carry on prescribed burning.

Attitudes Toward Prescribed Burning

Within the Wildlife Management

Profession

Primarily because of Stoddard’s influence, backed by his

rigorous studies of bobwhite quail, wildlife biologists in

the South had accepted prescribed burning, at least in

principle, before other resource management

professionals. The basic techniques of controlled

burning for game management were well established by

the mid-1930’s. The optimum season, frequency, and

conditions for burning for various game species and

important food plants were recognized. And special

techniques, such as spot-burning at night to create a

mosaic of vegetation, were in use (Stoddard 1935b).

Prescribed burning on an annual or biennial rotation

was a standard practice in the management of bobwhite

quail. In 1932 Stoddard reported “occasional controlled

burning of overly ‘rough’ cover” was among “measures

practiced on scores of Southeastern preserves. . .” for

wild turkeys (Stoddard 1932); he recommended specific

burning techniques on a 2- to 4-year rotation (Stoddard

1935a; 1939). And, although restoration of white-tailed

deer (Odocoileus virginianus) populations in the South

was just beginning, controlled burning was used to

improve browse conditions in areas of the Coastal Plain.

As previously noted, burning of marshes to improve

habitat for waterfowl and muskrats (Ondatra zibethica)

was in practice in the 1930’s or earlier (Lynch 1941).

Outside the South, wildlife biologists were skeptical or

more cautious. Despite research showing improved

habitat conditions for many species following

experimental and wild fires, prescribed burning was not

commonly employed for the management of wildlife.

With few exceptions, books on wildlife management

before 1960 treated prescribed burning as a practice

peculiar to the South. From 1930 through the 1950’s

there were seven important textbooks on wildlife

management. With one exception, all of them

emphasized protection from fire and treated prescribed

burning as a practice limited in application to the

southern coastal plain.

In his classic textbook “Game Management,” Aldo

Leopold (1933), who was well acquainted with

Stoddard’s work, made only passing mention, in a

scattered half dozen sentences, of controlled burning.

He referred to spot burning of heather for red grouse

(Lagopus lagopus) and to Stoddard’s use of fire in

bobwhite quail management to control nesting cover.

He mentioned Stoddard’s suggestion that controlled

burning may reduce ectoparasites and might prove

useful as a means of range sanitation, but then wrote

“Use of fire without sound evidence of its effects, or on

lands primarily devoted to other crops, is of course to be

deplored.”

In “The Land and Wildlife” Edward H. Graham (1947)

briefly acknowledged the use of fire in management of
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 17

Gulf Coast marshes for muskrats and snow geese (Chen

caerulescens). Regarding use of prescribed burning on

forest lands, he wrote only two sentences: “It has been

used in longleaf pine forests of Georgia to check growth

of hardwoods and increase wild legumes, a device for

improving habitat for bobwhite quail” and “although

fire is a tool sometimes employed by both foresters and

wildlife managers, its use is specialized and does not

negate the general rule of protection from burning.”

Reuben Trippensee (1948), in his widely used textbook,

referred to controlled burning as “a management

technique peculiar to the Southeast, where certain

vegetative cover types not only permit the use of fire in

this manner but apparently require it if quail range is to

be kept productive.” Citing Stoddard (1931, 1939), he

devoted nearly two pages to controlled burning, but

only for quail management in longleaf pine. Regarding

wild turkeys, he stated “in all regions except the

Southern Coastal Plain, lack of forest-fire control means

exclusion of the turkey.”

Wallace Grange’s book “The Way to Game Abundance”

(1949) provided the one notable exception in its

treatment of fire as a habitat management tool. Grange

considered animal cycles in the North to be largely a

result of plant succession linked to a cycle of drought

and associated fires. An entire chapter was devoted to

controlled burning for accomplishing a variety of

objectives in the management of various species. Grange

stated “with the possible exception of the desert, all

other North American game habitats have been, from

time to time, improved by the agency of naturally

occurring fire, and may be benefited through the

application of controlled fire.”

Leonard Wing’s textbook “Practice of Wildlife

Conservation”(1951) dealt similarly with the subject.

Wing made the statement that “The value of controlled

burning is becoming recognized, but a clear distinction

should be made between controlled and uncontrolled

burning.” He then devoted about a page to the practice

of controlled burning in the Southeast. He gave no

examples from other areas but did suggest that

controlled burning might be useful in the management

of some other game birds and specifically mentioned

songbird and non-game conservation.

As late as 1959 Ira Gabrielson, first director of the U.S.

Fish and Wildlife Service and long-time president of the

Wildlife Management Institute, in his textbook “Wildlife

Conservation” mentioned prescribed burning in a

paragraph devoted to Stoddard’s work with quail, then

added “Generally speaking, however, on the basis of

present knowledge, fire is so great an enemy of both

wildlife and forests that there is an increasing amount of

fire control, both on public and on private lands. This

should in the long run be exceedingly beneficial to

wildlife” (Gabrielson 1959).

Into the 1980’s most textbooks and references on

wildlife management continued to reflect little

awareness of or interest in prescribed burning.

Roy Komarek (1966) criticized wildlife professionals for

neglecting habitat management in general and

prescribed burning in particular. He noted that the index

to the first 29 volumes of the Journal of Wildlife

Management had no entries for “controlled burning” or

“prescribed burning”, and only seven entries to

“burning” or “fire.” He found the subject similarly

lacking in the other publications of the Wildlife Society.

Writing that the wildlife profession was lagging in the

use of fire, he called for more experimentation with

season, frequency, and size of prescribed fires for

different objectives and different species, including nongame species and wild flowers. Previously a strong critic

of the conservative attitude of foresters toward

controlled burning, he noted that by this time foresters

had perfected techniques by which they were burning

hundreds of thousands of acres. He wrote “Despite the

fact that Stoddard pioneered the use of controlled

burning on game lands more than thirty years ago, the

field of wildlife management, except in a few cases, has

contributed little to the advancement of the art and in

many cases depends upon techniques developed by the

foresters.”

Continuing Stoddard’s promotion of the study of fire

ecology and the controlled use of fire in land

management, the Tall Timbers Research Station was

formed in 1958, and through its research,

demonstration plots, and especially its annual fire

ecology conferences and published proceedings,

contributed greatly to the study, understanding and use

of fire. Stoddard, brothers E. V. and Roy Komarek, and

others at Tall Timbers deserve much credit for the

acceptance of prescribed burning outside the Deep

South. At the first North American Wildlife Conference,

Stoddard (1936) had urged experimentation with

prescribed burning in other regions. And, from the

beginning, the fire conferences were worldwide in scope,

bringing in speakers from all continents to address the

ecology and use of fire in many vegetation types and

parts of the world, emphasizing that the usefulness of

prescribed burning was not limited to management of

longleaf pine.

Prescribed Burning in Nongame

Wildlife Management

Although management of nongame wildlife is a

relatively new emphasis, recognition of the value of fire

as a potential tool for nongame wildlife management is

not. Most prescribed burning has multiple objectives,

and just because prescribed burning was not conducted

specifically for nongame objectives does not necessarily

mean that managers were unaware of the effects on

nongame species and did not take them into
18 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

consideration when burning. In his early writings,

Stoddard, who was an esteemed ornithologist, made

frequent references to effects of fire on species other

than game, forage, and commercially important timber.

In one of his early publications (Stoddard 1936) he

wrote that “multitudes of small birds” use the burns in

late winter and early spring and “such burns (are)

important to their welfare also.” In one of his reports to

cooperators, he included a section titled “burning to

maintain floral beauty of pinelands” (Stoddard 1939).

Stoddard’s recommendations for burning in quail

management included discussion of fire’s effects on

rodent and predator populations and its role in reducing

predation on quail. E. V. Komarek, working with

Stoddard, elaborated on the effects of controlled

burning on mammals in a paper for the Journal of

Mammalogy (Komarek 1939). Komarek’s later writings

included frequent references to the benefits of fire to

wildflowers, earthworms, butterflies, and birds

(Komarek 1969, 1971).

But, until the 1970’s there was little emphasis in the

South or elsewhere in managing specifically for

nongame wildlife (Landers and Johnson 1980), so

nongame species were only rarely a specific objective of

prescribed burning. Interest in managing habitats for

species other than game animals greatly increased in the

1970’s. This was evident in the surge in conferences,

symposia and special sessions at technical meetings

devoted to nongame species (DeGraaf 1978; Odom and

Guthrie 1981; Odom and Landers 1978; Thompson

1971). Although most effort in nongame and

endangered species management at that time was

devoted to determining the status of these species,

prescribed burning was considered in papers making

recommendations for habitat management for various

reptiles and amphibians (Herman 1981; Landers and

Speake 1980; Means and Campbell 1981; Means and

Moler 1978), nongame birds (Edwards 1978; Meyers

and Johnson 1978; several papers in Thompson 1971;

Wood and Niles 1978) and mammals (Hilliard 1979;

Lustig and Flyger 1975; Winchester et al. 1978).

Also, it was recognized that prescribed burning for

forestry or game management objectives maintained a

biotic community with a distinct suite of plant and

animal species (Komarek 1971; Landers and Speake

1980), and prescribed burning received increasing

attention for community restoration and maintenance

in parks and natural areas (Hendrickson 1972).

We end this historical review with a symposium on

prescribed fire and wildlife in southern forests held at

Myrtle Beach, South Carolina in 1981 (Wood 1981). The

symposium included papers on the status of prescribed

burning programs on state, federal, and private lands

and a series of papers reviewing the database on the

effects of prescribed fire on many individual species or

groups of vertebrates. The presentations demonstrated

that, although there was generally strong conceptual

understanding of fire ecology and its relation to habitat

needs of various wildlife and there was a good database

for some game species, with a few exceptions, data on

fire and non-game species were seriously inadequate.

Implications

The most obvious lesson of the controlled burning

controversy in the first half of the Twentieth Century

should be a reminder that all knowledge does not reside

with professionals or government agencies. In 1981 E. V.

Komarek noted that private hunting preserves of the

Southeast for more than 50 years had burned between

750,000 and 900,000 acres annually, leading the way in

spite of intense discouragement by government agencies

and many resource management professionals

(Komarek 1981).

Secondly, this review shows that conditions are always

changing. Fire has been a significant environmental

factor throughout much of North America for thousands

of years. But its role has been a dynamic one—

determined by climatic shifts and for several thousand

years by the ever-changing human factor. There is much

interest now in managing for natural landscapes. But, it

is not possible to define what was the natural condition,

except by defining the exact place and time—and

accepting human influences as natural. Managers could

set a goal of re-creating conditions of the sixteenth

century when the Spanish explorers passed through the

South. But, early descriptions of the landscape are spotty

and probably biased; and, even if early conditions were

known, they might not be what is desired today. Early

accounts indicate that Indians used fire excessively in

many areas, and the landscape they created in some

areas may have excluded forest cover that would be

valued today. Furthermore, in most places it would not

be possible to re-create prehistoric conditions because

soils have been drastically changed, important species

have been lost, and exotics have become established.

Therefore, we think prescribed burning strategies should

be based on objectives that are better defined than vague

notions of what is natural.

Third, land managers must be able to adapt to

continuing change. With increasing human populations

and urban-suburban encroachment on rural areas, there

will be increasing concerns about public health,

automobile accidents, and liability resulting from

problems with air quality and smoke produced by

prescribed burning. These concerns may result in serious

limitations on the use of prescribed burning in the

future. And, if, as many scientists contend, the climate is

becoming warmer and drier, this may drastically alter

the use of fire—for example, making hazard reduction a

more important objective and more seriously restricting

the season of burning. Also, history shows that priorities

in resource management are controlled by economic

and political conditions. Controlled burning conducted

by herdsmen 100 years ago was well suited to their
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 19

objective; but with different economic conditions (i.e.,

development of the forest industry), changes were

needed. Today, much prescribed burning has multiple

objectives, even in wildlife management. Current

interest in management for game and nongame wildlife

and natural areas may wane if economic prosperity

declines or there is an urgent need for other forest

products. Therefore, development of techniques for a

diversity of approaches to fire management is needed to

allow compromise and adaptability in meeting future

needs. Research also must continue on alternative

methods of vegetation control because the use of

prescribed burning will almost certainly become

restricted in some areas.

Finally, fire research is still needed. Managers have been

able to draw on 300 years of burning experience and

more than 50 years of fire research in the South. But,

since the 1981 symposium on prescribed fire and

wildlife, prescribed burning has been applied to an

increasing variety of situations where new objectives are

in place. Management objectives include an increasing

variety of plant and animal species and communities.

Prescribed burning is no longer considered to be out of

bounds of consideration in the management of any

habitat type. It is being used, at least in a limited way, in

the mountains for site preparation, hazard reduction,

oak regeneration, maintenance of bog turtle (Clemmys

muhlenbergi) habitat, maintenance of grassy balds as

habitat for small mammals and raptors, and

management of plants of special interest. Growing

season fires are being used increasingly for hardwood

control, promotion of flowering of grasses and other

forbs, and to produce diversified patterns of plant

growth. Infrequent high intensity fires are being used to

restore or regenerate communities, and new technology

for igniting and controlling fires is being developed by

foresters. With more diverse objectives and increased

use of prescribed burning in habitats other than

southern pine forests, the need for research is greater

than ever, and the research database may be weaker in

relation to needs than it was in 1981, when the

prescribed fire and wildlife conference was held. Roy

Komarek’s criticisms of the wildlife profession in 1966

remain valid today. We think this is especially true

outside the South.

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24 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Abstract.—Experimentally evaluating the success of

hardwood reduction techniques against a “model”

reference condition of longleaf pine sandhill

communities is not directly possible because reference

sites are not randomized or replicated. We addressed

this issue by measuring the similarity of arthropods in

treatment (fire, herbicide, felling/girdling, and control)

and reference sites using three indices. Arthropod

assemblages from plots that burned were significantly

more similar to those in the reference condition than

were those that did not receive fire. Our findings suggest

that increasing arthropod prey using fire will be

beneficial both to management and to biodiversity.

Introduction

Southern longleaf pine (Pinus palustris P. Mill.) forest

restoration is, in many ways, essentially synonymous

with red-cockaded woodpecker (Picoides borealis Vieillot)

habitat management, where mechanical removal and

short burn rotations (1–3 years) are used to reduce and

control midstory hardwood encroachment that is

detrimental to the woodpeckers (Conner and Rudolph

1989; Hooper et al. 1991). Because many public lands

in the southern U.S. are now managed for red-cockaded

woodpeckers (Brennan et al. 1995; U.S. Forest Service

1995), management effects on non-target species,

especially birds, have become a concern in recent years

(Brennan et al. 1995; Burger et al. 1998; Hunter et al.

1994; Plentovich et al. 1998; White et al. 1999).

Red-cockaded woodpecker habitat management (i.e.,

fire) likely affects arthropods directly or indirectly by

stimulating population growth through increased

resource availability and quality (James et al. 1997; Reed

1997). It is well documented that fire causes plants to

resprout and to be more palatable to herbivores (smaller

C:N ratio for plant tissues) (Dunwiddie 1991; Nagel

1973; Owensby et al. 1970; Smith and Young 1959;

Stein et al. 1992). Increased flowering after fire (Platt et

al. 1988) should attract insect pollinators. In turn,

increases in herbivores and pollinators may attract

predatory arthropods and parasites. Some predatory

ground beetles, for example, increase following fire

(Harris and Whitcomb 1974). Increases in arthropods

are important because they represent a fundamental

prey base for red-cockaded woodpeckers and associated

vertebrates, many of which exclusively feed arthropods

to their offspring and depend on arthropods for food

outside of the breeding season. Conversely, these species

of concern may suffer if fire suppression results in

decreases in arthropod abundance. Remarkably, there is

a dearth of quantitative data published on the

relationship between fire and the arthropods that redcockaded woodpeckers consume (but see Hanula and

Franzreb 1998; James et al. 1997) or on arthropods in

longleaf pine forests (reviewed in Folkerts et al. 1993),

especially describing old-growth communities.

We experimentally compared the effects of three

hardwood reduction techniques (growing season burn,

herbicide application, and midstory mechanical felling/

girdling) and a no-treatment control on herb-layer

arthropod densities in fire-suppressed longleaf pine

sandhills at Eglin Air Force Base (EAFB), Florida.

Hardwood reduction is necessary to meet EAFB’s goals

for restoring functional, diverse sandhill systems across

the Base and for habitat restoration for the endangered

red-cockaded woodpecker and other target species

having similar habitat requirements. Our main goal was

to identify which management technique(s) caused

arthropod species assemblages to converge toward

values found in fire-maintained sandhills, which we

termed the “reference condition” to emphasize forest

composition and function in addition to structure.

Based on our analysis, we also developed metrics that

may be indicators of sandhill management success.

Methods

Site Description

Eglin Air Force Base (EAFB) occupies 187,555 ha in the

southern portions of Santa Rosa, Okaloosa, and Walton

counties in the western Florida Panhandle. The mean

annual temperature is 18.3° C, with approximately 275

freeze-free days per year. Mean annual precipitation is

158 cm. Monthly precipitation levels peak slightly

during late spring and early summer months and

decrease during the winter months (Chen and Gerber

1990). Since 1886, 25 tropical storms and 27 hurricanes

made landfall within 97 km (60 miles) of EAFB.

The terrain is level to gently rolling with occasional areas

of steep slopes along creeks. Elevation ranges from 0–

100 m above sea level, and the landscape generally

slopes to the southwest toward the Gulf of Mexico.

Lakeland series, the common soil, is a medium to fine

sand with 5-10 percent silt and clay (Overing et al.

Fire, Herbicide, and Chainsaw Felling Effects on Arthropods in Fire-Suppressed

Longleaf Pine Sandhills at Eglin Air Force Base, Florida

Louis Provencher1, Krista E. M. Galley1, Andrea R. Litt1, Doria R. Gordon2,

Leonard A. Brennan3, George W. Tanner4, and Jeffrey L. Hardesty2

1

The Nature Conservancy, Niceville, FL 32588

2 The Nature Conservancy, Gainesville, FL 32611

3Tall Timbers Research Station, Tallahassee, FL 32312

4

Department of Wildlife Ecology and Conservation,

University of Florida, Gainesville, FL 32611
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 25

1995), is rapidly permeable and strongly acidic, with

nearly level to steep slopes. The vegetation of EAFB’s

sandhills is described in Rodgers and Provencher

(1999).

Restoration Experiment: Experimental

Hardwood Removal Plots

A total of 24, 81-ha plots were established in six blocks

of four fire-suppressed hardwood-longleaf pine sandhill

plots across EAFB (see map in Rodgers and Provencher

1999). This study utilized a randomized complete block

design (Steel and Torrie 1980), where each plot within

an experimental block was randomly assigned to either

control designation (no treatment), or to one of three

restoration treatments applied during the spring and

early summer of 1995: growing season burn in May or

June, herbicide application (ULW®

, the granular form of

hexazinone with 75 percent active ingredient applied at

2.24 kg/ha), and oaks and sand pine felling/girdling by

chainsaw (slash not removed). Fuel reduction burns

were conducted in the herbicide and felling/girdling

plots in early 1997. In each restoration plot, 32, 10  ́ 40-

m subplots were located in the 20-ha corner farthest

from the neighboring plots of the block to reduce the

potential for recording organisms that can travel across

adjacent plot boundaries (Figure 1).

Reference Plots

Three pairs of 81-ha frequently burned, longleaf pinedominated reference sandhill plots were established

(Fig. 1). These plots were not part of the experiment

described above but were used to measure restoration

success. In each reference plot, 32, 10  ́ 40-m subplots

were located in the center of the plot to reduce edge

effects (Figure 1). These plots are more fully described in

Rodgers and Provencher (1999) and Provencher et al.

(2000; 2001a; 2001b). For the duration of the

restoration study, reference plots were under a “let burn”

management policy (all reference plots burned once and

four plots burned at least twice during the study period).

Data Collection

Densities of selected herb-layer arthropod species were

estimated. To successfully collect arthropods of various

sizes and mobility, individuals were first collected using

a sweep net, followed immediately by a modified D-Vac

insect vacuum. In 1994, individuals were collected from

Figure 1.—Sample layout of one of six reference plots (81-ha) and of hardwood

reduction plots and sampling areas in one of six blocks in a randomized complete

block split-plot design consisting of four whole-plot treatments. Spatial randomization

of treatments varies per block.

450 m

GROWING

SEASON BURN 

GRANULAR

HEXAZINONE

(ULW®)

900 m

CHAINSAW

FELLING / GIRDLING 

900 m

CONTROL

(NO TREATMENT) 

240 m

Sub-Subplot Sampling Layouts Within 10 × 40-m Subplot 40 m

10 m

Reference Subplot Layout

240 m

Hardwood Reduction Plot & Subplot Layout

Four corner invertebrate (pre-treatment) sub-subplots: 0.5 × 2 m. One central

quantitative invertebrate sub-subplot added post-treatment: 0.5 × 8 m. 
26 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

herb-strata vegetation (<1.4 m) within four 0.5  ́ 2-m

sub-subplots (Figure 1). Because we suspected that the

noise and motion of our vacuum device and sweep net

were flushing some arthropods from adjacent subsubplots, we changed the location and shape of

arthropod sampling sub-subplots to a single 0.5  ́ 8-m

rectangle in the center of the subplot beginning in fall

1995 (Figure 1). Moreover, the second method

minimized escapes by arthropods because we opened

the sweep net once instead of four times. Specimens

from sweep net/D-Vac samples were manually sorted

and preserved in 70 percent ethanol. We enlisted the

assistance of taxonomic specialists, where possible, to

perform initial species identifications and established a

reference collection of more than 300 species of

authoritatively identified adult arthropods.

Statistical Analyses

Similarity measures allowed us to directly determine

which treatment plots most resembled the reference

plots. Treatment differences were tested using two-way

analysis of covariance (ANCOVA). The advantages of

this method over discriminant function analysis were

that (1) we could include both common variables and

those with patchy distributions and (2) the calculated

similarity values, rather than the raw data, were required

to meet any statistical assumptions of the tests.

Although there are a wide variety of available similarity

indices, we calculated similarity of selected variables

between each treatment plot and each reference site

using three formulas (Brower et al. 1989; Underwood

and Chapman 1998). The purpose of using three indices

was to achieve a consensus among them, especially for

the identification of indicator variables (see below). The

strength of an indicator increased with the number of

indices identifying it as a significant contributor to the

pattern of similarity.

Proportional similarity (PS) was selected because it is

widely used and it accounts for the relative abundance

of variables. This last feature was especially important to

us because site conditions (e.g., soil productivity) may

greatly change the values of certain variables but not

their relative abundance on a particular sampling site.

The PS between each treatment plot i (= 1,..., 24) and

each reference plot j (= 1,...,6) was averaged over all

reference plots per restoration plot i with a sum

weighted by sample sizes;

PSij = 

j =1

6

∑ nj

 (1–0.5

k=1

K

∑ |pik–pjk|)/N

where pik is the proportion of the logarithm of variable k

in treatment plot i, pjk is the proportion of the logarithm

of variable k in reference plot j, nj

 is the number of

subplots in reference plot j, and N is the total number of

subplots in all reference plots (Brower et al. 1989). We

took the logarithm of variables to prevent large values

from dominating PS and, thus, to increase the

representation of uncommon species. Plots that share all

the same variables in the same proportions will have a

PS = 1, whereas plots that share no variables will have a

PS = 0.

Following Underwood and Chapman’s (1998)

description of its superior properties, we calculated the

weighted average of the 1 – Bray-Curtis index of

dissimilarity,

BCi

 =

j =1

6

∑ nj

(1–

k=1

K

∑ |Zik–Zjk|/(∑ ∑ = =

+

N

k

N

k 1

jk

1

Zik Z ))/N

where Zik is the mean abundance of species k in plot i.

This index does not require proportional abundance or

any transformation and is, therefore, more intuitive than

proportional similarity, but it is interpreted in the same

way as proportional similarity.

We created a third similarity index, endpoint difference

(ED), bounded by 0 and 1 to incorporate within-plot

variation,

EDi

 =

j =1

6

∑ nj

k=1

K

∑ exp[–|Zik–Zjk|/σeij]/K/N

where the value within the exponential function is the

absolute value of the t statistic (Steel and Torrie 1980),

seij is the joint standard error of Zik and Zjk assuming

unequal sample sizes, K is the number of variables, and

Zik is either the average of a single variable (no

logarithmic transformation) k from plot i (similarly for

plot j) or the average proportional value of variable pik.

of variable k. Plots that share the same variables in the

same proportions (all |Zik–Zjk| = 0) or that have large

variability (standard errors are large) preventing these

plots from being differentiated will have an ED equal or

close to 1. Plots that share no variables (all |Zik–Zjk| are

large) or that can be easily distinguished due to low

variability (standard errors approach 0) will have an ED

close to 0.

To determine which variables contributed most to the

similarity pattern among treatments, we correlated the

24 similarity values with the contribution of each

species. A positive correlation would indicate that the

variable supported the similarity pattern, whereas a

negative correlation would mean that the variable

weakened the similarity pattern. We retained only

variables with significant correlations (> 0.481 or < –

0.482, df = 23, P < 0.05) (Steel and Torrie 1980). For

proportional similarity, the contribution of each variable

per plot i was arbitrarily measured by:

∑=

6

j 1

(1–0.5|pik–pjk|)/6

where we averaged contributions over the six reference

sites per variable. Similar averaging and calculations

were performed for the 1 - Bray-Curtis and endpoint

difference contributions:
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 27

∑=

6

j 1

(1–|Zik–Zjk|/(∑ ∑ = =

+

N

k

N

k 1

jk

1

Zik Z ))/6, and

∑=

6

j 1

exp[–|Zik–Zjk|/σeij]/6, respectively.

We tested restoration treatment effectiveness by

comparing the similarity indices using two-way

ANCOVA for a randomized complete block design (Steel

and Torrie 1980). Pre-treatment similarity was used as

the covariate to account for differences among plots that

existed prior to treatment application. Keeping within

the maximum number of allowable independent

contrasts for three degrees of freedom (Sokal and Rohlf

1981), we contrasted the following treatments: control

versus spring burn (C vs. B), burn versus the herbicide

ULW®

 (B vs. H), and herbicide versus felling/girdling (H

vs. F) (see Provencher et al. 2000b for more

information). Similarity values were transformed with

x , x2

, or log(x) to stabilize variances when necessary.

Results and Discussion

Similarity to the Reference Condition

Significant increases in similarity for herb-layer

arthropods between spring burn and reference plots

were not observed until fall 1996, although endpoint

difference, while congruent with the other indices, was

not significantly affected by treatments (Fig. 2A). The

fact that arthropod similarity was only responding to

fire was instructive. The arthropods that we sampled

appeared to be closely tied to the plants as resources, but

not to the percentage of hardwood reduction. In 1996,

one year after initial treatment, felling/girdling topkilled

the most oaks (93 percent), followed by herbicide (69

percent), and fire (18 percent), compared to firesuppressed control plots, where the highest oak

densities were found (1330 stems/ha ± 60) (Provencher

et al. 2001b). On the other hand, fire causes

groundcover plants to resprout, providing tender,

nutritious forage promoting the growth of plant-eating

populations (e.g., grasshoppers, leafhoppers,

planthoppers) or groups that feed on live and dead

vegetation (e.g., springtails) (Dunwiddie 1991; Nagel

1973; Owensby et al. 1970; Smith and Young 1959;

Stein et al. 1992). Herbivorous arthropods and

springtails are among the numerically dominant species

we sampled, therefore strongly affecting similarity. In

turn, a greater availability of arthropods lower in the

food web should attract predators and parasites such as

spiders, wasps, and some flies and beetles. Indeed, the

parasitic (braconid) wasps Chelonus sp. and Heterospilus

spp. and the spiders Tmarus rubromaculatus (Keyserling),

Misumenops spp., Mimetus sp., and Hentzia palmarum

(Hentz) were among the 20 more common species in

burn and reference plots during the fall 1996.

Fuel reduction burns applied to herbicide and felling/

girdling plots during the winter and spring of 1997

presented several opportunities to test if the response of

herb-layer arthropods in herbicide and felling/girdling

plots would follow the same pattern observed in the

spring burn plots. We suspected that the effect would be

stronger as plant resprouting was expected to be more

complete (due to the continuous fuel matrix created by

oak leaf drop and felled oaks). Burning should result in

approximately 50 percent topkill and abundant

resprouting (Glitzenstein et al. 1995), while herbicide

application should practically eliminate oaks and

significantly reduce resprouting.

Spring 1997 was a rather unusual field season as it

immediately followed fuel reduction burns. As a result,

similarity analyses were potentially influenced by half of

the plots being charred. This does not offer a compelling

test of the hypothesis. Not surprisingly, the spring burn

plots were closest to the reference condition for all

indices, but only significantly so for proportional

similarity and 1 – Bray-Curtis (Figure 2B).

Arthropod data collected in Fall 1997 showed the

strongest convergence to that in the reference condition.

Despite the hot fires in felling/girdling-fire plots, the

herb-layer arthropod assemblage was more similar to

the reference assemblages in the fire only and felling/

girdling-fire than in the herbicide-fire plots for all

indices (Figure 2B). We attribute the difference between

the herbicide and felling/girdling treatments to the

greater herbaceous (mostly grasses) cover in the former

treatment, a resource that herbivores are expected to

exploit. The fire-suppressed control plots were still the

least similar to the reference plots. The hypothesis that

arthropods would rapidly benefit from the more

thorough coverage of the hotter fires was confirmed:

arthropods responded within 6 months after the 1997

fires, whereas a response came 1 full year after the spring

burns of 1995. Seasonality of fire should not explain

this effect because most fuel reduction burns were

completed in the early spring. As in 1996, the reduction

of hardwoods was not a factor that explained similarity

results, the greatest total cumulative topkill of oaks

occurred in the herbicide-fire plots (94 percent), with

lower mortality in the felling/girdling-fire plots (62

percent) and the prescribed fire only plots (41 percent, a

delayed mortality relative to 1996) (Provencher et al.

2001).

Although we hypothesized that the growth of arthropod

populations benefited from newly resprouting palatable

vegetation, by 1997 this explanation no longer applied

to the plots burned two years earlier. This implies that

once established, arthropod populations persist beyond

when plant palatability is a key factor and/or the

continued resprouting of hardwoods (Provencher et al.

2001a) is utilized as a source of palatable vegetation.

Our data do not allow evaluation of the duration of fire

suppression necessary before insect populations again

become depressed.
28 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Figure 2A.—Time series of three indices (proportional similarity, 1 –

Bray-Curtis index, and endpoint difference) measuring the similarity of

herb-layer arthropods between each of four hardwood reduction

treatments (spring burn, herbicide, felling/girdling, and no-treatment

control) and reference plots from fall 1994-1996 at Eglin Air Force Base,

Florida. Tests of treatment effects were calculated with two-way ANCOVA.

The experimental design is a randomized complete block (6 blocks),

split-plot design, but only the block design at the whole plot level is

presented here. The covariate was the pre-treatment data from the fall

1994 or spring 1995. The error term is the mean square of the interaction

of the block and restoration treatment effects. Similarity values received

various transformations when it was necessary to stabilize variances. The

center of the box is the mean, the edges of the box correspond to one

standard error, and the error bars are a 95 percent confidence interval.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 29

Figure 2B.—Time series of three indices (proportional similarity, 1 –

Bray-Curtis index, and endpoint difference) measuring the similarity of

herb-layer arthropods between each of four hardwood reduction

treatments (spring burn, herbicide, felling/girdling, and no-treatment

control) and reference plots from spring 1997-1999 at Eglin Air Force

Base, Florida. Tests of treatment effects were calculated with two-way

ANCOVA. The experimental design is a randomized complete block (6

blocks), split-plot design, but only the block design at the whole plot

level is presented here. The covariate was the pre-treatment data from the

fall 1994 or spring 1995. The error term is the mean square of the

interaction of the block and restoration treatment effects. Similarity

values received various transformations when it was necessary to stabilize

variances. The center of the box is the mean, the edges of the box

correspond to one standard error, and the error bars are a 95 percent

confidence interval.
30 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

The similarity patterns seen in 1997 were repeated in

spring and fall 1998 and spring 1999, with increased

convergence toward the reference condition to some

limited extent for herb-layer arthropods in herbicide

plots (Figure 2B). Felling/girdling plots showed the

same response to a lesser extent (Figure 2B). Fire appears

to be the cause of these changes.

Indicators of restoration success

Two valuable outcomes of identifying the species or

groups that contributed most to the similarly evaluation

(indicator) are to reveal ecological relationships and to

identify candidate variables that may be included in a

monitoring program. We include as candidate indicators

those variables that significantly contributed to the

similarity of at least two of the three similarity indices.

Selection of arthropod indicators was difficult because

of the great difference among similarity indices. For

example, we never identified more than three species

with the Bray-Curtis-based index, whereas >10 species

were identified by the other indices. The springtail

Sminthurus sp. 1, the leafhoppers Erythroneura spp., the

planthopper Metcalfa pruinosa (Say), the leafhopper

Jikradia olitoria (Say), and the jumping spider H.

palmarum were the strongest indicators and the

dominant species to increase after spring burning or

immediately after the fuel reduction burns (Fig. 3).

Except for the spider, these arthropods are herbivores or

detritivores (the diet of Sminthurus sp. 1 is unknown but

herbivory is suspected). Earlier we explained the

relationship between fire, palatable plant resprouts, and

arthropod population growth. These indicators illustrate

this scenario well. These spiders are generalist predators

and may themselves be tracking arthropod prey

densities. (Although not evident in Figure 3 for fall

1996, average H. palmarum densities were higher in the

burn plots than other treatments only after logarithmic

transformation.)

During fall 1997, the first time the three similarity

measures showed significant treatment differences,

Sminthurus sp. 1 and the leafhoppers Empoasca spp.,

another common herbivore, were the strongest

contributors to all the indices. The jumping spiders

Habronattus spp. were also positively and significantly

correlated to the similarity pattern, but they were

uncommon in our samples. Endpoint difference was the

only index that did not differentiate among the

treatments in spring 1998, therefore suggesting that high

variance in species density affected the treatment

comparisons. The common leafhopper J. olitoria was the

sole indicator shared by proportional similarity and the

Bray-Curtis-based index, thus replacing Sminthurus sp. 1,

which was not a strong contributor to any of the indices.

All three indices showed significant treatment

differences in the following two seasons. In fall 1998,

the leafhoppers (Empoasca spp.) were the group to

contribute significantly to more than one index

(proportional similarity and endpoint difference). These

species’ response to all hardwood reduction methods,

especially fire, was pronounced as virtually no

individuals were ever recorded from control plots

(Figure 3). As in the previous season, no species

significantly influenced all three indices during spring

1999. Sminthurus sp. 1, the jumping spider #25

(unidentified), the planthopper Oecleus sp., and the ant

Crematogaster ashmeadi Mayr were the only indicators

identified by at least two indices. Because the restoration

effect for endpoint difference was barely significant (P =

0.046), jumping spider #25, Oecleus sp., and C. ashmeadi

may not be the most reliable indicators of restoration

success. This left Sminthurus sp. 1 as the strongest

potential indicator, achieving its highest densities in

felling/girdling plots, somewhat lower in herbicide and

burn plots, and the lowest densities in control plots

(Figure 3).

Overall, very few species dominated the list that we

would suggest as indicators of restoration success over

time: Sminthurus sp. 1, J. olitoria, Empoasca spp.,

Erythroneura spp., and M. pruinosa. With the exception of

Empoasca spp., these species were generally common and

statistical analyses could be performed on their densities

(see Figure 3). Clearly, there was a connection between

fire and the increased densities of these arthropods: we

hypothesize that the palatability of resprouting plants is

the main cause of this interaction.

Management Implications

• Long-term management of sandhills and other

communities on Eglin on a large scale requires

economical methods that stimulate productivity in

the understory. Only fire will satisfy these two

constraints for herb-layer arthropods.

Prescribed burning is substantially cheaper to apply

than herbicide and felling/girdling. In 2000, burns

started with ground ignition cost approximately $21.90/

ha, whereas aerial ignition can now be accomplished for

$8.60/ha (J. Furman, EAFB, pers. comm.). Herbicide

application operations amount to $140.80/ha for the

herbicide and $76.57/ha for labor (total $217.40/ha).

Chainsaw felling/girdling is $158.10/ha, involving only

labor. These estimates do not include the cost of a fuel

reduction burn that would normally follow herbicide

and felling/girdling operations.

• The relationship between fire, understory arthropods,

and the diet of red-cockaded woodpeckers requires

further research as this connection affects their

management and that of other species in longleaf pine

forests.

The diet of red-cockaded woodpeckers during the

breeding season depends exclusively on arthropods.

Hanula and Franzreb (1998) have shown that 70
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 31

Figure 3.—The densities of herb-layer arthropods identified as the strongest indicators of

restoration success by similarity analyses in hardwood reduction and reference plots at

Eglin Air Force Base, Florida, from fall 1996 to spring 1999. Indicators were only selected

when similarity values were significantly different among hardwood reduction treatments.

Tests of treatment effects were calculated with two-way ANCOVA. The experimental design

is a randomized complete block (6 blocks), split-plot design, but only the block design at

the whole plot level is presented here. The covariate was the pre-treatment data from the

fall 1994 or spring 1995. The error term is the mean square of the interaction of the block

and restoration treatment effects. Density values received various transformations when it

was necessary to stabilize variances. The center of the box is the mean, the edges of the box

correspond to one standard error, and the error bars are a 95 percent confidence interval.

C B H F R

4

3

2

1

0

Erythroneura spp.

C B H F R

0.35

0.30

0.25

0.20

0.15

0.10

0.05

0

–0.05

Hentzia palmarum

C B H F R

0.30

0.22

0.14

0.06

–0.02

Fall 1996

Spring 1997

3

2

1

0

Metcalfa pruinosa

2.0

1.5

1.0

0.5

0

Fall 1997

1.2

1.0

0.8

0.6

0.4

0.2

0

–0.2

35

30

25

20

15

10

5

0

1.6

1.2

0.8

0.4

0

–0.4

C B H+B F+B R

P = 0.0686

a

b

c

c

Not tested

P = 0.0189

Sminthurus sp. 1 Empoasca spp.

Sminthurus sp. 1 Jikradia olitoria

Sminthurus sp. 1

P = 0.1732

a

b

c c

P = 0.0001

P = 0.8278

a

b

c c

P = 0.0189

a

b

c c

P = 0.0072

Spring 1998

Fall 1998

Jikradia olitoria

1.6

1.2

0.8

0.4

0

Empoasca spp.

0.4

0.3

0.2

0.1

0

Crematogaster ashmeadi

C B H+B F+B R

0.4

0.3

0.2

0.1

0

Sminthurus sp. 1

C B H+B F+B R

6.5

5.5

4.5

3.5

2.5

1.5

0.5

–0.5

Spring 1999

P = 0.0073

a

b b

b

P = 0.0510

a

b

b

b

P = 0.1400

Not tested

Density / 4 m2

Treatment Treatment Treatment

Treatment Treatment Treatment
32 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

percent of the arthropod prey consumed by redcockaded woodpeckers below the crown of longleaf

pines disperse from the understory (mostly ground/soil

arthropods). Therefore, there appears to be a strong

dietary link between fire, understory arthropod

population growth, and the woodpeckers’ reproductive

success (see also James et al. 1997).

• Little is known about arthropods in second- and oldgrowth longleaf pine forests. Quantitative and

taxonomic research on this subject would greatly help

both ecologists studying other aspects of longleaf pine

forest ecology and managers needing to conserve

arthropod communities and recover the populations

of threatened and endangered species.

We identified potential indicators to monitor that may

inform managers on longleaf pine sandhill quality and

fire intervals. It is worthwhile to determine if decreasing

similarity of herb-layer arthropods between managed

and reference sites may roughly correspond with the end

of a fire-free interval.

Acknowledgments

We thank the staff of the Jackson Guard Natural

Resources Management Division at Eglin Air Force Base

for providing the financial resources to conduct this

research. We are also grateful for additional funding

from The Nature Conservancy’s Florida Regional Office.

K. Campbell, A. Iosue, E. Levine, S. McAdoo, J. McAdoo,

M. Northrup, and L. West helped with data collection

and entry. This effort was sponsored by the U.S. Army

Medical Research and Material Command, Department

of the Army under Cooperative Agreement number

DAMD17-98-2-8006.

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hardwood reduction techniques in fire-suppressed

sandhills in northwest Florida. Forest Ecology and

Management. 148: 63-77.

Provencher, L.; Herring, B. J.; Gordon, D. R.; Rodgers, H.

L.; Galley, K. E. M.; Tanner, G. W.; Hardesty, J. L.;

Brennan, L. A. 2001b. Effects of hardwood reduction

techniques on longleaf pine sandhill vegetation in

northwest Florida. Restoration Ecology. 9: 13-27.

Reed, C. C. 1997. Responses of prairie insects and

other arthropods to prescription burns. Natural

Areas Journal. 17: 380-385.

Rodgers, H. L.; Provencher, L. 1999. Analysis of longleaf

pine sandhill vegetation in northwest Florida.

Castanea. 64: 138-162.

Sokal, R. R., and F. J. Rohlf. 1981. Biometry, 2nd edition.

New York, New York: W. H. Freeman and Company.

859 p.

Smith, E. F.; Young, V. A. 1959. The effect of burning on

the chemical composition of little bluestem.

Journal of Range Management. 12: 139-140.

Steel, R. G. D.; Torrie, J. H. 1980. Principles and

Procedures of Statistics, 2nd edition. New York, New

York: McGraw-Hill Book Company. 633 p.

Stein, S. J.; Price, P. W.; Abrahamson, W. G.; Sacchi, C. F.

1992. The effect of fire on stimulating willow

regrowth and subsequent attack by grasshoppers

and elk. Oikos. 65: 190-196.

Underwood, A. J.; Chapman, M. G. 1998. A method for

analyzing spatial scales of variation in composition

of assemblages. Oecologia. 117: 570-578

U.S. Forest Service. 1995. Final Environmental Impact

Statement for the management of the red-cockaded

woodpecker and its habitat on national forests in

the southern region. Management Bulletin R8-MB

73. Atlanta, GA: U.S. Department of Agriculture,

Forest Service, Southern Region.

White, D. H.; Chapman, B. R.; Brunjes, IV, J. H.;

Raftovich, Jr., R. V.; Seginak, J. T. 1999. Abundance

and reproduction of songbirds in burned and

unburned pine forests of the Georgia Piedmont.

Journal of Field Ornithology. 70: 414-424.
34 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Abstract.—In western North America, major wildfires

often now result in stand-replacement events and

natural resources losses for many decades post-burn.

Fire severity has been exacerbated by past fire

suppression that has allowed large fuel load

accumulations. To reduce woody debris and restore the

ecological integrity of western forests, prescribed

burning is increasingly used as a regional management

tool. However, we do not understand the effects of

either wildfire or prescribed fires on amphibians in

stream, riparian and terrestrial habitats in western

forests. Terrestrial amphibians, macroinvertebrates and

other animals are surface active during periods of

rainfall or high moisture. Wildland fire usually starts in

the hot, dry summers typical of these more arid Western

and Mediterranean climates and may have less effect on

resident biota than prescribed fires often conducted

during the late fall to spring rainy season, when there is

sufficient moisture to prevent crown fires. Still, intense

wildfires may result in increased erosion and sediment

or changes in soil chemistry impacting downstream

aquatic environments. To our knowledge, no published

reports exist on effects of fire on the aquatic

herpetofauna of the Pacific Northwest. Research efforts

now underway include new studies of wildland fires in

Oregon and Idaho on aquatic amphibians, and studies

on the effects of prescribed fire on terrestrial

salamanders and associated forests in the Klamath

Province along the Oregon-California border. These will

help evaluate the cumulative effects of fuels reduction

on amphibian population and habitat structure, and

provide guidelines to better manage for wildlife species

characteristic of western forests. In the Pacific

Northwest, investigations of fire effects on wildlife are

severely lacking relative to the vast acreage, economic

value, and biodiversity of its forest ecosystems. Given the

increasing prominence of wildfire and prescribed

burning in many western forest systems, we suggest

more resources will be devoted to such research

endeavors, and that they include other sensitive groups

of wildlife such as mollusks.

Fire is a natural, recurring disturbance in forested

ecosystems of western North America, but it has been

aggressively suppressed for >50 years. Although fire

prevention was implemented to protect forest resources,

these efforts have resulted in greatly increased fuel levels

and, in turn, increased risk of catastrophic fires (Pyne

1982, Agee 1988, Henjum et al. 1994). Catastrophic

fires are generally defined as stand-replacing fires that

burn at spatial scales and intensities atypical to the

historic fire regime. For example, forested environments

with long fire return intervals (300-600+ years) are

subject to large-scale stand replacing fires because fuel

loadings are typically high (Morrison and Swanson

1990). Of greater importance might be how past forest

management activities such as clearcutting and thinning

and associated forest fragmentation affects the resulting

forest structure and the fire mosaic when they do burn.

In contrast, forests with historically short fire return

intervals (20-40 years) were characterized by reduced

fuel loads where stand replacing fires were less common.

Fire suppression is hypothesized to have the greatest

affect on forests with short (i.e., <50 year) fire return

intervals.

Resource managers now recognize the importance of fire

for maintaining healthy forests, yet often face conflicting

priorities when pursuing multiple resource management

objectives. Prescribed burning, prescribed wildland fire

(e.g., allowing natural fires to burn within specified

parameters), and other fuels management practices are

being introduced into the landscape to reduce fuel

loadings, but little is known about their effects on the

biota of the forests that are being restored or altered

(Potter and Kessell 1980). For example, we lack

information on the quantity and quality of the resulting

downed woody debris that comprises the critical habitat

for many species of resident wildlife. Current standards

and guidelines of the Northwest Forest Plan require

Federal land managers to promote retention of deadand-down wood as wildlife habitat yet reduce high fuel

loads through prescribed burning or other fuels

reduction practices (USDA/USDI 1994).

Further, we lack information on both the short- and

long-term suitability of the post-burn habitat for

resident and migratory wildlife. Fuels management will

alter the structure and composition of existing firesuppressed systems. Large wood plays a vital role in

ecosystem processes such as nutrient and water cycling

(Harmon et al. 1986, Franklin and Spies 1991). It

provides a moist, thermally stable habitat for many

species of wildlife both large and small (Maser and

Trappe 1984, Carey and Johnson 1995, Bull et al. 1997).

We lack basic information on the response of large

woody debris to burning and the suitability of firescarred woody debris for wildlife.

In Pacific Northwest forests, terrestrial salamanders are

strongly associated with structural elements of the forest

Responses of Amphibians to Fire Disturbance in Pacific Northwest Forests:

a Review

R. Bruce Bury1, Donald J. Major2 and David Pilliod3

1

USGS Biological Resources Division, Corvallis, OR 97331

2Department of Fisheries and Wildlife, Logan, UT 84321

3Aldo Leopold Wilderness Research Institute, University of

Montana, Missoula, MT 59807
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 35

floor such as woody debris, moss, and surface/subsurface rock (Bury and Corn 1988, Corn and Bury1990,

Welsh and Lind 1995). Stream amphibians also are

dependent on the structural components of stream

substrate including in-channel large woody debris (Corn

and Bury 1989). However, almost nothing is known

about the responses of amphibians and their habitats to

fire and fuels management practices in the West

(DeMaynadier and Hunter 1995, Welsh and Droege

2001). Only a few unpublished studies exist on the

effects of fire on pond-breeding amphibians in western

forests exist (Askey and Peterson 1993). These limited

data sets indicate no significant differences in

amphibian occurrence in ponds located in burned

versus unburned forests.

There are some recent studies on the effects of

prescribed fire on herpetofauna in the eastern U.S.

(Russell et al. 1999, Ford et al. 1999), where fire

frequency has been reduced and the season of burn

changed from mostly growing-season to dormantseason in Southeastern pine forests (Means and

Campbell 1980, Robbins and Myers 1992). Widespread

public and agency concerns over large fires are now

driving policy-level decisions to integrate fire and other

fuel reduction strategies into management of

ecosystems.

Our goal is to review and compare the effects of fire and

fuel reduction management on Pacific Northwest

amphibians and their associated habitats, especially

downed woody material. We will attempt to summarize:

(1) what we know generally from a few available studies,

and (2) what information is needed.

Stream Amphibians

There are three families (10 species) of amphibians

endemic to the rocky, cool-water streams in the

mountainous regions of the Pacific Northwest. Many of

these species are of conservation concern due to

declining populations (Bury 1994, Smith et al. 1998).

Therefore, understanding fire effects on stream

amphibians and their habitats will be of utmost

importance in future management practices in

Northwest forests. Some stream amphibians such as the

giant salamander (Dicamptodon spp.) may respond

positively to increased stream productivity (increased

periphyton or algae growth) for a few years after forest

opening from timber harvest (Murphy and Hall 1981)

and presumably fire. Stream amphibians also are

sensitive to changes in debris flow, elevated temperature,

water chemistry, and sedimentation that often occur

post-disturbance (Bury 1988, Gamradt and Kats 1997,

Kerby and Kats 1998, Welsh and Ollivier 1998). For

example, the tailed frogs (Ascaphus spp.) are among the

least tolerant of the anurans to elevated temperature and

will die quickly when exposed to water temperatures at

or near 29.6°C (deVlaming and Bury 1970). In contrast,

giant salamanders tend to have broader tolerance to

temperature and siltation than other stream amphibians

(Bury 1988).

Although direct mortality of aquatic and wetlandassociated life stages of amphibians may be low where

wet areas and riparian zones provide refugia from fire

(Vogl 1973), terrestrial life stages in nearby uplands may

experience much higher mortality associated with the

direct and indirect effects of fire that alter prey

availability or change shelter and microclimate (Lyon et

al. 1978, 2000; Russell et al. 1999). Further, physical

and biological changes in adjacent uplands may

influence the survival and well-being of biota in riparian

zones through downslope effects on hydrology and

water chemistry in streams (Minshall 1989).

Some research on large, stand-replacing wildfires on

lotic biota suggests that fire ultimately benefits aquatic

invertebrates and fishes, even those species that are

negatively affected by the disturbance immediately after

the fire (Roby and Azuma 1995, Minshall et al. 1997,

Rieman et al. 1997). For example, Lyon et al. (1978)

suggested that some aquatic invertebrates may decline

immediately after a fire, then increase to levels above

pre-fire conditions as a response to increased stream

productivity. Large fires can have long-term effects on

streams by: (1) reducing invertebrate diversity for a

decade or longer (Roby and Azuma 1995); (2) changes

in peak discharge, stream channel morphology, large

woody debris inputs, and sediment loadings; and (3)

elevated temperature and altered water chemistry

(Richards and Minshall, 1992; Minshall et al. 1997).

Also, the effects of fire on stream biota may be more

pronounced in headwater streams than in mid-order or

larger streams (Roby and Azuma 1995, Minshall et al.

1989, Minshall et al. 1997).

Although there is little empirical evidence of how fire

impacts stream amphibians, we can glean some

information from the Old-growth Wildlife Habitat

Program in the Pacific Northwest (Ruggerio et al. 1991),

which compared biota across chronosequences of forest

categorized into young, mature, and old-growth stands.

Because almost all of these stands were naturally

regenerated from wildfires, the younger stands were

most recently burned. Thus, fire-sensitive wildlife

populations would be expected to be different in

younger stands compared to older, mature stands that

had a longer time since disturbance.

Spies (1991) estimated the age of trees across the

chronosequence of stands in the Oregon Coast Range

(Table 1) and reported that the young and mature

stands were not the equivalent of intensively-managed

plantations resulting from harvest. For example, there

often were large amounts of downed woody debris in

young natural stands, resulting from input of fallen trees

that were fire killed. There was less wood volume in

mature stands and then large accumulations again in

old-growth. Managed stands tended to have low
36 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

amounts of downed wood once slash rotted or was

burned on site. Clearcut stands lack large trees and, in

turn, recruitment of large downed woody material on

the forest floor.

Bury et al. (1991a) found no significant difference

among stream amphibians across natural young,

mature, and old-growth stands in three Northwest

biogeographic provinces (Table 2). However, Corn and

Bury (1989) reported that the density and biomass of

four species of stream-associated amphibians were

significantly greater (2-7X) in uncut forest (natural

regeneration) than in streams in clearcut stands sampled

14-40 yr post harvest. Although there were few young

natural stands (n = 3), the abundance of amphibians

was similar to streams found in mature and old-growth

stands (Table 3), but had markedly greater numbers

compared to streams running through clearcut stands.

This evidence suggests that wildland fire had little effect

on stream amphibians or amphibians recovered rapidly

in postburn conditions, but clearcut logging was

detrimental and had long lasting effects. However,

further study is needed on this topic, especially with

more replicates of study streams.

Studies on prescribed and wildland fire effects on

stream-breeding amphibians in the Northwest are

currently underway. In Idaho, we (Pilliod and P. S.

Corn) have initiated a 3-year study comparing Rocky

Mountain tailed frog (Ascaphus montanus) populations

in streams running through both burned and unburned

forests. In conjunction with this retrospective study, we

are conducting an experimental prescribed fire study in

the South Fork Salmon sub-basin where Rocky

Mountain tailed frog and Idaho Giant salamander

(Dicamptodon aterrimus) populations will be monitored

in six streams for 3 years pre- and 2 years post-burn.

Similar research will start in Oregon in 2002 by one of

us (Bury).

Riparian Habitats

Riparian zones adjacent to streams are important

habitats for Pacific Northwest amphibians (Bury 1988).

Most of the endemic stream amphibians of the region

inhabit the waters and adjacent cool, vegetated banks.

Riparian vegetation serves to shade and cool stream

temperatures, and protect the rocky substrate from

siltation from upslope disturbance. Stream amphibians

like tailed frogs (Ascaphus truei) often move into the

riparian zone after transformation (T. Wahbe, pers.

comm.), where conditions are relatively cool and moist

year-round compared to hotter, drier upslope areas.

In forested stands west of the Cascade crest, riparian

areas associated with permanent streams are not

scheduled for prescribed burning (T. Atzet and J. Lint,

pers. comm.). These Federal lands have been protected

as riparian reserves under the Northwest Forest Plan

(NWFP). The Aquatic Conservation Strategy of the

NWFP provides riparian reserves of 1-2 tree heights away

from all permanent streams. In 1997, an unpublished

NWFP report on “Riparian Reserve Evaluation of

Techniques and Censuses: Federal Guide for Watershed

Analysis” suggested that management activities in

riparian areas are feasible, including fuels reduction.

Introduction of prescribed fire into riparian areas

appears to be only allowable on an experimental basis

for research or pilot studies at this time. However,

prescribed fire likely will occur near riparian zones and

there may be an influence from these adjacent, upslope

areas.

Fire suppression in riparian areas for long periods will

create fuel build-ups that may eventually lead to

unnaturally severe fire in riparian zones, especially in

the more productive sites at lower elevations. Further,

disruption of the natural fire regime likely will result in

changes in riparian forest structure and composition.

This may include increased amounts of input of woody

debris into streams that, in turn, influence channel

processes and habitat availability for stream or

streamside amphibians.

Terrestrial Habitats

To our knowledge, there are no studies addressing the

effects of fires on terrestrial amphibians in the Pacific

Northwest. However, we can again glean useful

information from the Old-growth Wildlife Habitat

Program (Ruggerio et al. 1991). Bury et al. (1991b)

summarized results for 130 stands in the

chronosequence from three Provinces (OR Cascades, OR

Coast Range, WA Cascades). Each stand was sampled in

the fall for one month of pitfall trapping (1,080 trapnights/stand), and most were sampled twice (1983 and

1984). This intensive effort showed that there were no

significant differences in catch of 6 of 8 common species

of amphibians along the chronosequence with young

stands did not differing from mature or old-growth. The

Northwestern salamander (Ambystoma gracile) was more

common in old-growth whereas red-backed

salamanders (Plethodon vehiculum) were more frequent

in young stands than in other types. However, both of

these species may be influenced by factors other than

stand age. The Northwestern salamander is a migratory

species that travels overland and location of breeding

ponds may greatly influence its local abundance. The

red-backed salamander requires rocky substrate and this

may have influenced its distribution. Overall, the

Table 1.—Estimated ages of trees in stands in the

Oregon Coast Range (from Spies 1991).

Stand type Young Mature Old-growth

Mean age 55 100 315

Range 30-79 84-120 130-525
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 37

resident amphibians were either little influenced by fire

or had recovered since burning 3-5 decades earlier.

Using pitfall trapping in the Oregon Coast Range, Corn

and Bury (1991) found no marked differences in

numbers of the five most common amphibians across

the chronosequence of natural regenerated stands (from

earlier fires): young, 40-75 yrs old (n = 8 stands);

mature, 80-120 (n = 10); and old-growth, 150-450 (n =

27). They also sampled 5 clearcut harvested stands (< 10

yrs post harvest) and overall abundance did not differ

much from results in natural stands (young to oldgrowth) except that tailed frogs and torrent salamander

(Rhyacotriton spp.) were absent. Both groups appear

associated with streams and both are highly sensitive to

logging.

Employing time-constrained searches of downed woody

debris, Corn and Bury (1989) found a correlation

between ensatina (Ensatina eschscholtzi) and clouded

salamander (Aneides ferreus) numbers and stand

amounts of downed wood. Loss of large wood input was

hypothesized to be a limiting factor for several species of

terrestrial plethodontids in Northwestern coniferous

forests.

In northern California, Welsh and Lind (1991) reported

that more species and numbers of individuals of

amphibians, especially terrestrial salamanders, were in

older forests compared to younger stands. These also

were in the naturally regenerated chronosequence. These

authors and others (Bury 1983, 1994; Smith et al. 1999;

Welsh and Droege 2001) suggest that timber harvest has

negative effects on several species of terrestrial

salamanders in western coniferous forests. Similarly,

there appear to be reduced numbers of salamanders in

logged stands in deciduous and mixed forests in the

eastern U.S. (Pough et al. 1987, Petranka et al. 1993,

deMaynadier and Hunter 1995). Timber harvest,

especially clearcutting, opens up forest canopies and

leads to desiccation. Removal of trees reduces input of

large chunks of woody debris into the forest ecosystem.

Further, harvested areas oftern are often subjected to

secondary site preparation or other pre-commercial

treatments such as burning, herbicide spraying and

thinning to enhance new tree growth. Although we

know little about how such multiple stressors work in

concert, timber harvest (especially clearcutting) and

associated silvicultural practices appear detrimental to

terrestrial amphibian populations (De Maydanier and

Hunter 1995, Welsh and Droege 2001).

To address how fire influences forest wildlife, we (Bury

and Major) are involved with a study of fire effects on

terrestrial amphibians in the Klamath Province

(southern Oregon and northern California). Key

objectives are to:

Table 3.—Stream-associated amphibians taken from three categories of forest stands

in the Oregon Coast Range. Results are mean numbers caught/10 m (most were

headwaters about 1 m wide).

Species Old-growth and Young Young

Mature Natural Harvest

(n = 20) (n = 3) (n = 20)

Tailed frog, Ascaphus truei 9.8 8.3 3.5

Torrent salamander, Rhyacotriton variegatus 3.1 1.7 0.4

Pacific giant sal., Dicamptodon tenebrosus 23.4 34.3 4.6

Dunn’s salamander, Plethodon dunni 4.6 3.0 1.4

Table 2.—Distribution of 79 streams by stand category and natural regenerated or

harvested state.

Natural Regeneration Clearcut Harvest

Young Mature Old-growth

Washington Cascades 6 6 6 0

Oregon Cascade 6 6 6 0

Oregon Coast Range 3 10 10 20
38 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

(1) Compare structural components of the forest

floor and their use by terrestrial herpetofauna in

burned and unburned sites;

(2) Determine the vulnerability of the structural

components used by herpetofauna; and

(3) Evaluate habitat quality by relating diversity

and abundance of herpetofauna to available forest

floor structure.

Our study has two designs: retrospective (wildland fire)

and experimental (prescribed fire with pre- and posttreatment). Retrospective work will describe forest floor

structure in burned and adjacent unburned sites in

recent (< 10 yr) wildland fires. We will conduct timeconstrained terrestrial surveys at the paired sites to

compare species richness and relative abundance of

herpetofauna, and to characterize the use of the

structural components by amphibians.

Our preliminary results from one large wildland fire in

the North Umpqua River Basin in Oregon in 1996

indicate that there was no negative effect from this fire

on terrestrial herpetofauna. We found more individuals

in the burned than unburned comparison areas using 3

paired plots. However, much more cover was available in

the unburned stands, so catch per cover object is

somewhat less in the burned forest. Lack of response by

the resident terrestrial amphibians may be related to

occurrence of this wildland fire in summer during

normally dry, hot conditions when terrestrial species of

amphibians are deep underground. However, lack of

information regarding pre-wildfire conditions (i.e., fuel

loadings, cover availability, and associated amphibian

detection rates) limits our inference capabilities. We

have no data on possible long-term effects.

Unlike most wildland fires, prescribed burning activities

often coincide with seasonal surface activity periods of

terrestrial salamanders in the spring and fall. We (Major

and Bury) are currently examining the effects of

prescribed fire on herpetofauna in late seral coniferous

and mixed-coniferous forests in the Klamath Province.

Specifically, we are examining pre- and post-burn

population-level responses of terrestrial salamanders

across a series of prescribed fires. We also will compare

structural components of the forest floor such as fire

fuels and salamander habitat. Finally, we hope to

explore the effectiveness of modeling population-level

responses of terrestrial salamanders to changes in forest

floor “habitats” through the use of predictive fuels

consumption and fire effects modeling. Field sampling

began in Summer 2001 with project completion

expected in 2003.

Discussion

The lack of information on the effects of fire on fish and

wildlife is a major impediment to developing

ecologically sound fire management policies (McMahon

and deCalesta 1990). Recent reviews of the effects of fire

on amphibian communities uncovered relatively few

studies, although most of the work reviewed had

occurred in Southeastern pine plantations (Russell et al.

1999). Most aquatic studies have focused on standreplacement burns and few have included an

experimental component, such as prescribed burns

(Major and Bury 2000). Further, many recent studies

have examined population descriptors such as species

presence or relative abundance, whereas more specific

information on population change and measures of

productivity may be needed to understanding causeand-effect relationships of fire in the Pacific Northwest.

Although recent directives promote fuels reduction, we

lack information on the quality of resulting postburn

habitats or how resident wildlife species respond to fireinduced changes in availability of the altered structural

elements (e.g., less forest duff, fewer cover objects).

Reduction in quantity of fuels, downed woody debris,

also could have a profound effect on the habitat needs

of wildlife and associated biological integrity that

federal land management is charged to conserve.

Overall, scientific studies on fire effects on wildlife and

associated habitats are just starting in the Pacific

Northwest. The current studies will provide some

information for evaluating the effects of fire and forest

management practices on several amphibian species in

stream, riparian, and terrestrial situations.

We believe there is a need for more investigations that

relate fuels management to habitat quality for resident

terrestrial herpetofauna for several reasons. Many

terrestrial amphibians are sensitive to habitat change

because they have moist, permeable skin and restricted

home ranges. Most terrestrial amphibians and some

reptiles also require large woody debris as nesting

habitat or cover objects (Welsh and Droege 2001). Thus,

herpetofauna are well suited as resident wildlife to

measure responses to habitat changes.

There is a need to assess prescribed fires inside riparian

reserves and adjacent slopes. For example, unburned

riparian areas likely buffer the stream from the effects of

fire immediately after the burn, but streams may have a

delayed response until spring runoff carries sediment

and nutrients into the stream during periods of peak

discharge. On longer time scales, excluding burning in

riparian areas may result in increased riparian forest and

amounts of instream woody debris dynamics. This may

increase amount of woody debris and pose greater fire

risk in riparian areas, especially during drought periods.

Amphibians may still respond to upland burning,

because juveniles and adults of stream amphibians

move into adjacent woods at the onset of fall rains and

fire-associated mortality of terrestrial phases of these

species may occur in prescribed fire periods.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 39

The few studies of fire effects on streams to date have

been conducted within single drainages or fire

complexes and have been focused on stand-replacing

wildfires. No published studies exist that address the

relative effects of stand-replacing and prescribed fire at

broad spatial scales. There is need for studies that

combine an ongoing prescribed fire study with post-fire

investigations of stream response to widespread

wildland fires. These will provide opportunities to

characterize the effects of fire treatments and severity on

aquatic and riparian habitats.

We need integrated regional and national programs that

better link wildlife habitat components into fuels

inventories and fire effects models. Although inventory

techniques are designed to collect information on

structural components pertinent to fuels management

goals, this information is of limited use in evaluating

“habitat use” by wildlife. Further, a habitat/fuels

approach would prove biologically useful in developing

effective monitoring protocols and providing baseline

information to elucidate working hypotheses on fire

effects and ecological responses across different spatial

scales. In the western U.S., much work has focused on

inventorying volume and tonnage of downed woody

fuels (Koski and Fischer 1979, Brown et. al. 1982).

However, these studies address only a subset of

structural components and do not provide information

on fire-mediated changes in quantity, type, size, and

physical characteristics of downed wood, which affects

wildlife habitat quality (Bull et. al. 1997). Fire effects

models (Peterson and Ryan 1986, Ryan and Reinhardt

1988, Keane et al. 1994) incorporate a wider range of

fuel types and more precise description of post-burn

conditions; however, they are limited in their utility to

describe habitat quality across the range of structural

legacy in northwest forests. Furthermore, these models

only examine immediate post-fire changes and are based

on modeling structural components influenced by past

suppression.

Clearly, we should increase numbers and integration of

projects that: 1) Determine what type and amount of

surface and ground structure are required by resident

wildlife, 2) Evaluate direct and indirect effects of fuels

management on structural components important to

resident wildlife, and 3) Predict the short- and long-term

effects of fuels management on legacy components in

forests and associated quantity and quality of habitat for

resident wildlife. There is a rich literature and growing

understanding of habitat requirements of forest wildlife,

in part related to major efforts generated by the

Northwest Forest Plan and other agency mandates. A

future goal is to tie this information to fuels inventory

models and integration of disciplines. Lastly, we need to

recognize the vast acreage and diversity of forests in

western North America, Geographically dispersed

studies will be required to better define how wildland

and prescribed fires affect wildlife across a broad

spectrum of environmental conditions.

Acknowledgments

We wish to thank E. J. Hyde, C. A. Pearl and K. Russell

for comments and discussion about this paper and

topic.

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The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 43

Abstract.—We examined the effects of rotational

livestock grazing and prescribed winter burning on the

state threatened Texas horned lizard, Phrynosoma

cornutum, by comparing home range sizes, survival

estimates and prey abundance across burning and

grazing treatments in southern Texas. Adult lizards were

fitted with backpacks carrying radio transmitters and

relocated daily. Prey abundance (harvester ants,

Pogonomyrmex rugosus) and activity were greater in

burned pastures, but grazing had a variable effect

depending on the timing since the last burn. Home

ranges in burned pastures were smaller than in

unburned pastures in the active season. Level of grazing

(heavy vs. moderate) did not affect home range size.

Summer survival rates of horned lizards were higher in

the moderately grazed sites than the heavily grazed sites.

The smaller home ranges, lack of effect on survival rates,

and greater prey abundance in burned pastures

suggested a positive effect of fire on Texas horned

lizards.

Introduction

The effect of land-use practices on sensitive species, such

as threatened or endangered species, is of considerable

conservation and political interest. However, little

information is available to evaluate the ecological effects

of management practices such as burning or grazing on

herpetofauna in general (Russell et al. 1999), and on the

threatened Texas horned lizard, Phrynosoma cornutum, in

particular. The Texas horned lizard is the official state

reptile of Texas (Donaldson et al. 1994), and is a species

of special concern in the conservation community.

Although the Texas horned lizard was protected by Texas

legislative mandate in 1967, it has declined throughout

its range, especially in Texas (Price 1990). Suggested

reasons for this decline include habitat alteration for

land uses such as agriculture or development, the

introduction of the red imported fire ant (Solenopsis

invicta), and the use of insecticides (Price 1990;

Donaldson et al. 1994). Such declines can decrease

genetic variability and hinder the lizard’s ability to

adjust to changes in environmental conditions caused

by land-use practices.

The habitat and prey of horned lizards can potentially

improve or worsen with fire and grazing. Fire reduces

shrub canopy cover (Dunne et al. 1991) where grazing

can increase the amount of woody vegetation (Archer

and Smeins 1991). Ruthven et al. (2000) found that

forbs increased on southern Texas rangelands in the first

year after a winter burn, but were not affected by

grazing. Bunting and Wright (1977) also found that

forbs and grass cover increased following fire. Ants, the

main prey of horned lizards, are not deleteriously

affected by fire or grazing (McCoy and Kaiser 1990;

Heske and Campbell 1991; McClaran and Van Devender

1995:165 Fox et al. 1996).

Specific objectives of our research were to: compare the

relative abundance and activity of harvester ants,

Pogonomyrmex rugosus, the main food source of the Texas

horned lizard, among different burning and grazing

treatments; and compare home range size and survival

rates of Texas horned lizards among different burning

and grazing treatments. Based on available literature, we

made several testable predictions. Harvester ant activity

and abundance should be greatest in the moderately

grazed/burned site because of an increase in seed

production coupled with an open, sparsely vegetated

habitat, selected by harvester ants (DeMers 1993).

Horned lizards will be less selective of foraging habitat

in the moderately grazed and burned site due to greater

prey abundance and better habitat interspersion.

Therefore, we predicted range size of Texas horned

lizards would be smaller and survival rates higher in the

moderately grazed/burned sites than in other

treatments.

Study Area

The 6,150-ha Chaparral Wildlife Management Area

(CWMA) occurs in Dimmit and La Salle Counties, Texas.

The CWMA was purchased by the state in 1969 and

management authority was given to the Texas Parks and

Wildlife Department (TPWD). Average annual rainfall

on CWMA is 63 cm with a primary peak in May and a

secondary peak in late September/early October (TPWD,

unpublished data). The dominant vegetation types on

the CWMA are honey mesquite (Prosopis glandulosa)

woodlands or parklands, with prickly pear cactus

(Opuntia engelmannii), tasajillo (Opuntia leptocaulis),

brasil (Condalia hookeri), spiny hackberry (Celtis pallida),

blackbrush acacia (Acacia rigidula), twisted acacia (Acacia

schaffneri), hogplum (Colubrina texenis), and Texas

persimmon (Diospyros texana) as common

subdominants. Common and scientific names for

vegetation follows Hatch et al. (1990).

Five study sites (50-60 ha) were selected on the CWMA,

each with a different burning and grazing treatment.

The Effects of Burning and Grazing on Survival, Home Range,

and Prey Dynamics of the Texas Horned Lizard in a Thornscrub Ecosystem

Anna L. Burrow1, Richard T. Kazmaier1, Eric C. Hellgren1, and Donald C. Ruthven, III2

1

Department of Zoology, Oklahoma State University,

Stillwater, OK 74078

2

Texas Parks and Wildlife Department, Artesia Wells, TX

78001
44 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Sites were chosen based on similarities in dominant

woody species and canopy coverages. Treatments were:

control (non-burned/non-grazed), moderately grazed/

burned, heavily grazed/burned, moderately grazed/nonburned, and heavily grazed/ burned. The control site has

not been burned or grazed since 1976.

Historical grazing occurred on CWMA, but after TPWD

began managing the land, grazing steadily declined and

temporarily stopped in 1984 because of poor range

condition. During this time, the grazing system was

changed from continuous grazing to different rotation

systems. Grazing resumed in 1991 with a high-intensity,

low-frequency rest-rotational system from 1 October to

30 April. Moderately grazed areas were stocked at 25

animal-unit days (AUD) • ha-1 • yr-1 and heavily grazed

areas were stocked at 37.5 - 50 AUD • ha-1 • yr-1. We

defined one AUD as 2 steers for one day.

A prescribed burning program was initiated on the

CWMA in 1997. Burns were conducted using head fires

ignited with a drip torch and covered 40 to 80 ha. The

study areas used in this research project were burned in

February 1998 and November 1999 during dry

conditions.

Methods

Field Methods

Lizards were captured in each of the study areas through

road cruising, fortuitous encounters, and drift fence

arrays. Each study site (n=5) on the study area had 3, Yshaped drift fence arrays that were open for 14 days in

either May or June. Upon capture, snout-vent length

(SVL), total length, mass, sex, and location of the lizards

were recorded. Lizards were marked with an intraabdominal passive integrated transponder (PIT; AVID,

Norco, California, USA) tag. The fifth toe on the front

right foot was also clipped to recognize if the lizard had

been previously caught. Lizards that were too small to

receive a PIT tag, approximately < 50 mm SVL, were

given a unique toe clipping sequence.

Adult lizards captured within the five study sites were

fitted with custom-made backpacks that carried

transmitters (150-151 MHz, L and L Electronics,

Mahomet, Illinois, USA). Backpacks were composed of a

beige muslin material and elastic straps dyed to match

the natural substrate color of the CWMA. The backpack

was attached to the lizard by placing an upper strap

around the neck and one front leg, and placing an

additional strap around the back legs. A drop of

cyanoacrelate gel adhesive was used to attach the straps

to the lizard’s chest and lower abdomen to further

secure the backpack. The total mass of the transmitter

and backpack bundle was approximately 3 g (< 8

percent of lizard mass). Receiving range of the

transmitters was approximately 100 m. An antenna

attached to the end of a 5-m PVC pole increased

transmitter detectability to approximately 200 m.

Radio-fitted lizards were initially relocated twice daily

with a handheld two-element Yagi antenna until lizards

resumed normal ranging behavior. Monitoring was then

reduced to once daily until hibernation. Every six weeks,

lizards were recaptured and given a new transmitter in

the field. Once refitted with a backpack, the lizard was

released. Data recorded at each relocation included

lizard activity and behavior, date, time of day, pasture,

burn treatment, UTM coordinates, weather, and microhabitat data.

Data were collected during the summers of 1998-2000.

The summer was divided into 2 seasons, active and

inactive, corresponding to the relative activity of horned

lizards. The season encompassing 15 April - 30 June was

the time of greatest lizard activity and was considered

the active season. Lizards are considerably more

sedentary during 1 July - 15 August, which was termed

the inactive season.

Locations of lizards were estimated by pacing from the

position of the lizard in a cardinal direction to a road

and then to a permanent landmark with known

Universal Transverse Mercator (UTM) coordinates.

Coordinates were entered into a Geographical

Information System (GIS) to aid in range and habitat

analyses. Only lizards with a radiotransmitter were used

in home range and survival rate analyses (Munger

1986). All statistical analyses were considered significant

at a = 0.10.

Ant Abundance and Activity

Ant abundance and activity were measured with bait

stations composed of six petri dishes placed 15-m apart

along a transect. Transects were randomly located and

followed a compass bearing. Each bait station was

baited with millet and was anchored to the ground with

a nail to prevent rodents from removing the dishes. Four

transects from the same study area were run

simultaneously. Petri dishes were baited in the morning

and checked between 0800 and 1100 for ant activity to

encompass the peak activity of ants (Whitford and

Bryant 1979). Number of ants foraging at the station

and the number of ants visiting the station within one

minute were recorded. Though other species of ants

were noted if present, only harvester ants were counted.

The bait stations were baited again in the evening to

assure that ants would keep visiting the dishes. Bait

stations were run for 4 days at a time using the same

transects once in the active season and once in the

inactive season in all five study areas in the summers of

1999 and 2000. Systematic searches for ant mounds

(Whiting et al. 1993; Fair and Henke 1997) were not

used in this study because it was difficult to determine

harvester ant mounds from other ant mounds, and to
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 45

distinguish active ant mounds from inactive ant

mounds.

Harvester ant abundance was averaged across the 4 days

for each transect, providing 4 replicates for each study

site. Differences in ant activity and abundance were

compared across the different burning and grazing

treatments using a repeated measures, 4-way ANOVA

including burning (burn, unburn), grazing (moderate,

heavy), year (1999, 2000), and season (active, inactive)

as main effects and all interactions. The treatments

creating the repeated measurements were season and

year. Because of a small sample of lizards in the control

site (ungrazed, unburned), and the lack of an ungrazed,

burned site, the control site was not used in the ANOVA.

Instead, we examined the following contrasts: control vs.

grazed sites, control vs. the grazed, unburned sites, and

control vs. the heavily grazed, burned site (the most

disturbed site).

Home Ranges

Range size of lizards were calculated using 95 percent

minimum convex polygons (MCP; Mohr 1947) with the

Animal Movement Analysis Program (Hooge et al.

1999). We included lizards tracked for 3 20 locations to

ensure a reliable representation of the home range.

Because the home range data were not normally

distributed, home range sizes were log-transformed.

Individual lizards were used as the experimental unit,

although this represents pseudoreplication, because the

treatments were not replicated (Hurlbert 1984).

Therefore, any inferences made from these data should

be used with caution beyond the study area.

Comparisons of range size were made with a 3-way

ANOVA including burning, grazing, and season as main

effects and all interactions. Preliminary analysis

indicated no gender differences in range size. Therefore,

data were pooled across sex. Contrasts to the control

were calculated in the same manner as described above

in the ant analyses.

Survival

Survival rates were estimated using the Kaplan-Meier

procedure (Pollock et al. 1989). Only lizards tracked for

3 10 days were used in these analyses. The fate of many

lizards was unknown due to transmitter failure, removal

by a predator, lizard migration, and discovery of a

backpack (without a lizard). Lizards with an unknown

fate were termed censored in the analyses. To determine

survival, we estimated the fate for censored lizards based

on knowledge of that lizard. Lizards for which a fate

could not be estimated were considered alive. To test for

differences in the survival function (shape of the curve)

between treatments, a log-rank test was used (Pollock et

al. 1989). A Z-test statistic was also used to compare the

survival curves on the last day of the summer (August

15; Pollock et al. 1989).

Results

Ant Abundance and Activity

More harvester ants were found at the bait station in the

burned pastures, but this effect varied by season and

level of grazing (3-way interaction, F1,36 = 5.00, P = 0.03,

Table 1.). In both seasons, more ants were found in the

burned pastures than the unburned pastures; however,

this was especially true in the inactive season. More ants

were found in all sites in the inactive season. In the

active season, similar numbers of ants were found in the

burned sites. In the inactive season, more ants were

found in the moderately grazed/burned site than the

heavily grazed, burned site (P < 0.01). In both seasons,

the number of ants at the bait station was similar in the

unburned sites. More ants were found in the control

than the unburned, grazed sites (F1,75 = 7.66, P < 0.01).

Fewer ants were found in the control than the 4 grazed

sites (F1,75 = 15.72, P < 0.01) and the heavily grazed,

burned site (F1,75 = 12.24, P < 0.01, Figure 1).

Burning affected the number of ants that arrived at the

bait stations in one minute, but this effect varied by

season and year (3-way interaction, F1,36 = 3.30, P = 0.07,

Table 2). More ants were found in the burned sites in

both seasons and both years than the unburned sites (P

< 0.04 for all comparisons), except for the active season

of 1999 (P = 0.47). In both years, more ants were found

in the inactive season than active season for burned sites

(P < 0.01 for both comparisons). More ants were also

found in the burned sites in 2000 when compared to

1999 for both seasons (P < 0.01 for both comparisons).

Finally, the number of ants that visited the bait stations

was similar in the unburned sites for both seasons and

both years.

Grazing also affected the number of ants that arrived at

the bait station, but this effect varied by season and year

(3-way interaction, F1,36 = 7.55, P < 0.01, Table 1). In the

Table 1.—Number of ants at the bait station upon

arrival (n = 4 in each treatment) for the active and

inactive seasons on the Chaparral Wildlife

Management Area, summers 1999 and 2000. The

treatments are designated as U-U (control), Mg-B

(Moderately grazed, burned), Mg-U (Moderately

grazed, unburned, Hg-B (Heavily grazed, burned),

and Hg-U (Heavily grazed, unburned)

Active Season Inactive Season

Treatment × SE × SE

U-U 8.3 4.5 5.3 1.6

Mg-B 5.8 2.4 28.6 3.9

Mg-U 1.8 1.3 5.2 2.1

Hg-B 9.6 3.0 16.6 3.0

Hg-U 2.8 1.2 4.8 1.4
46 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

active season, the number of ants that visited the bait

stations was similar in the moderately and heavily

grazed sites. However, in the inactive season of 1999,

more ants were found in the moderately grazed sites

than the heavily grazed sites (P = 0.05); whereas in 2000

during the inactive season, more ants were found in the

heavily grazed sites than the moderately grazed sites (P <

0.01). In both years, more ants were found in the

inactive season than the active season for all sites (P <

0.01 for all comparisons) except for the heavily grazed

sites in 1999 (P = 0.51). More ants were found in 2000

than 1999 for both seasons and levels of grazing (P <

0.04 for all comparisons).

More ants arrived at the station in the control than for

the 4 grazed sites (F1,75 = 3.52, P = 0.06), but fewer than

in the heavily grazed, burned site (F1,75 = 5.86, P = 0.01,

Figure 1). More ants were found in the control than the

unburned, grazed sites, but this difference was not

significant (F1,75 = 1.23, P = 0.27). Though this

comparison was not tested, fewer ants were found in the

control than the burned, grazed sites (Figure 1).

Home Ranges

A total of 78 seasonal home ranges from 57 lizards were

used in home range analyses. Total area used by horned

lizards across both seasons ranged from 0.02 ha to 11.05

ha for 95 percent MCP (Table 3). The effect of burning

on home range size interacted with season for 95

percent MCP (F1,14 = 3.49, P = 0.08). In the active

season, home ranges in the burned sites (‘x ± SE = 1.14

± 0.27 ha, n = 18) were smaller than those in the

unburned sites (2.01 ± .06, n = 19), but were smaller

and similar in size in both sites during the inactive

season. All other interactions and main effects were not

significant. Grazing did not have an effect on home

range size (P = 0.15). Average (± SE) home range size for

lizards in the control was 0.66 (± 0.22) in the active

season and 0.80 (± 0.28) in the inactive season. None of

the contrasts to the control were different.

Survival

Summer (15 Apr - 15 Aug) survival rates (S) ranged

from 0.25 to 0.62. Grazing influenced survival rate of

lizards (P = 0.05, Figure 2). Survival rates of lizards in

the moderately grazed sites (S = 0.60) were higher than

those in the heavily grazed sites (S = 0.36). Burning did

not affect summer survival rates (P = 0.19, Figure 3).

Because of a small sample size in the control, lizards

from the control were not used in these analyses, but

summer survival rate in the control was 1.00. However,

2 of 4 lizards died in the September - October period.

Discussion

The diet of the horned lizard consists primarily of ants

(Burt 1928; Milne and Milne 1950; Pianka and Parker

1975; Whitford and Bryant 1979; Rissing 1981; Munger

1984a; Munger 1984b; Schmidt et al. 1989). Pianka and

Parker (1975) found that 69 percent of the diet of Texas

horned lizards was composed of harvester ants, with

beetles composing the remainder. Numbers of harvester

ants, therefore, could be one of the main components

determining optimal habitat for a horned lizard.

Previous studies on the effects of burning and grazing

on ants showed that ants are not deleteriously affected

by fire or grazing (McCoy and Kaiser 1990; Fox et al.

1996). The large increase in ant numbers for all ant

indices in burned sites compared to unburned sites

implies that on the CWMA, burning was beneficial to

harvester ants. This conclusion was supported by the

contrasts to the control. Ant numbers in the ungrazed,

unburned control were intermediate to low values on

the grazed, unburned sites, and high values on the

grazed, burned sites.

The interpretation that burning had a positive effect on

ant activity and distribution was complicated by

interacting effects with grazing, season, and year.

However, examination of these interactions indicated

that, with regard to burning, the treatment effect varied

Figure 1.—Contrasts to the control for the number of

ants at the ant bait stations upon arrival and the number

of ants that visited the station within one minute at the

Chaparral Wildlife Management Area, summer 1999-

2000. The treatments are designated as Mg-B

(moderately grazed, burned), Mg-U (Moderately grazed,

unburned), Hg-B (Heavily grazed, burned), and Hg-U

(Heavily grazed, unburned).

0

5

10

15

20

Ants at the Station

0

10

20

30

40

50

Control MG-B MG-U HG-B HG-U

Ants/Minute
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 47

Table 2.—Number of ants that visited the bait stations within one minute (n = 4 in each treatment) for

the active and inactive seasons on the Chaparral Wildlife Management Area, summers 1999 and 2000.

The treatments are designated as U-U (control), Mg-B (Moderately grazed, burned), Mg-U (Moderately

grazed, unburned, Hg-B (Heavily grazed, burned), and Hg-U (Heavily grazed, unburned)

1999 2000

Active Season Inactive Season Active Season Inactive Season

Treatment × SE × SE × SE × SE

U-U 1.3 1.0 7.7 2.9 35.5 10.7 40.4 12.9

Mg-B 3.1 1.5 45.7 12.8 28.4 73.7 73.7 11.3

Mg-U 0.1 < 0.1 5.8 2.5 12.8 6.7 10.2 5.1

Hg-B 9.4 4.9 15.2 5.1 33.4 8.4 101.8 17.3

Hg-U 1.2 0.2 5.6 2.2 17.5 5.4 39.1 7.2

Table 3.—Home range sizes of Texas horned lizards (ha) using 95% Minimum Convex Polygon (MCP) for active

and inactive seasons on the Chaparral Wildlife Management Area, summer 1998-2000. The treatments are

designated as U-U (control), Mg-B (Moderately grazed, burned), Mg-U (Moderately grazed, unburned), Hg-B

(Heavily grazed, burned), Hg-U (Heavily grazed, unburned)

Active Season Inactive Season

Treatment n × SE minimum maximum n × SE minimum maximum

U-U 6 0.66 0.22 0.06 1.28 4 0.80 0.28 0.22 2.11

Mg-B 8 1.04 0.50 0.04 4.26 10 0.32 0.10 0.02 0.96

Mg-U 9 1.33 0.20 0.25 2.11 5 0.23 0.05 0.10 0.35

Hg-B 10 1.22 0.31 0.03 3.13 6 0.80 0.28 0.22 2.11

Hg-U 10 2.62 1.02 0.06 11.05 10 0.49 0.02 0.04 1.93

0

0.2

0.4

0.6

0.8

1

0 25 50 75 100 125

Interval Day

Survival

M oderate (n = 26)

Heavy ( n =37)

15 Apr 18 Aug

P = 0.05

Figure 2.—Survival rates of Texas horned lizards in

the moderately grazed (S = 0.60, 95% CI = 0.37-

0.83) and heavily grazed sites (S = 0.36, 95% CI =

0.16-0.55) of the Chaparral Wildlife Management

Area, summers 1998-2000.

0

0.2

0.4

0.6

0.8

1

0 25 50 75 100 125

Interval Day

Survival

Burn (n = 32)

Unburn (n = 31)

15 Apr 18 Aug

P = 0.19

Figure 3.—Survival rates of Texas horned lizards in

the burned (S = 0.52, 95% CI = 0.30-0.73) and

unburned (S = 0.38, 95% CI = 0.16-0.60) sites of

the Chaparral Wildlife Management Area, summers

1998-2000.
48 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

only in magnitude by season and year. The direction of

the burning effect did not vary with time. Foraging

activity in Chihuahuan desert harvester ants varied with

season and year, with greater numbers of foragers in July

and August than May and June (Whitford and

Ettershank 1975). The greater number of harvester ants

on the CWMA in the inactive season could be a result of

increased foraging effort by harvester ants caused by

greater seed availability in the inactive season, coupled

with an increase in the number of foragers from

reproductive efforts in early summer (Whitford and

Ettershank 1975). Harvester ants are also thermophilic

(Holldobler and Wilson 1990), so it is possible that the

hotter temperatures associated with the inactive season

enable harvester ants to forage more.

Increased seed availability and reproduction can also

explain year effects. Whitford and Ettershank (1975)

stated that harvester ant activity was regulated by seed

availability and colony satiation. Perhaps 2000 was an

exceptionally good year for seed production or

reproduction resulting in more foraging ants. Increased

ant activity in 2000 also could be a result of depleted

resources from 1999. If granaries were depleted in 1999

due to a bad seed year, foraging effort would increase in

2000 to attempt to replenish the granaries. Low seed

production in 1999 would also result in reduced activity

in that year (Whitford and Ettershank 1975), thereby

reducing the number of foraging ants at the bait

stations. Unfortunately, we do not data on seed

production to support these speculations.

Previous studies concluded that livestock grazing did

not affect ant numbers in desert ecosystems (Heske and

Campbell 1991; McClaran and Van Devender

1995:165). Grazing had a variable effect on harvester

ant numbers, and appeared beneficial to ants when

coupled with burning (Figure 1.1). As with burning,

effects were stronger in the inactive season. The effect of

level of grazing on ant activity interacted with year and

specific ant index. For example, in the inactive seasons,

more ants arrived at the station in moderately grazed

sites in 1999, but in heavily grazed sites in 2000.

However, more bait stations were consistently visited by

ants in heavily grazed sites. In addition, higher ant

indices were seen on the heavily grazed/burned site in

all contrasts with the control. More bare ground in the

heavily grazed site could be responsible for the increase

in ants in heavily grazed sites in 2000, because harvester

ants prefer areas of sparse vegetation (Holldobler and

Wilson 1990). Therefore, it appears that heavy grazing

was more beneficial to harvester ants, especially when

coupled with burning.

Fire and grazing can improve conditions for harvester

ants in several ways. Fire can increase forb and grass

cover (McClaran and Van Devender 1995:134; Ruthven

et al. 2000) and available bare ground, and decrease

litter accumulation. Grazing can also increase forb

abundance and decrease litter accumulation (Kelting

1954). Because harvester ants are granivores, most

activity occurs in areas with interspersed bare ground

and herbaceous vegetation. Forb and grass seeds provide

the ants with food, and the sparse vegetation facilitates

foraging (Holldobler and Wilson 1990). Finally, DeMers

(1993) noted that harvester ant queens prefer to start a

new mound in open areas with little vegetation. An

increase in prey abundance associated with burning

could result in a subsequent increase in Texas horned

lizard density or a decrease in ranging behavior.

Home range size is inversely proportional to the

distribution and abundance of resources for many

species, including several lizards (Mares et al. 1976;

Litvaitis et al, 1986; Boutin 1990; Lacher and Mares

1996). Little is known about the size of home ranges for

the Texas horned lizard, although information does exist

in closely related species (Lowe and Stebbins 1954;

Baharav 1975; Turner and Medica 1982). Munger

(1984c) reported home range sizes of Texas horned

lizards in Arizona as averaging 1.35 ha for females (n =

13) and 2.40 ha for males (n = 10). Home ranges in my

study were considerably larger than those previously

reported by Fair and Henke (1999), who estimated

home range size of Texas horned lizards in southern

Texas to be between 0.02 to 1.47 ha (n = 16). However,

their home range estimates were only the sum of weekly

estimates of several months.

We propose that the smaller home ranges of horned

lizards in burned pastures resulted from burning

improving the habitat of horned lizards to the degree

that ecological requirements (i.e., food, cover) were

found in a smaller area. Because grazing did not affect

home range size, it is possible that grazing at the

intensities studied is neither beneficial nor harmful to

the habitat of horned lizards. The mechanism by which

burning and moderate grazing may improve horned

lizard resource distribution, and thus reduce home

range size, is by creating open habitats interspersed with

vegetation cover. Whiting et al. (1993) found that Texas

horned lizards selected disturbed habitats over

undisturbed habitats. They suggested that prey

abundance and suitable open habitats were major

factors related to the spatial occurrence of Texas horned

lizards in Texas. Disturbances that create an open,

sparsely vegetated habitat appear to benefit horned

lizards in several ways. Open areas facilitate movement

by this dorso-ventrally flattened species (Whiting et al.

1993). Fair and Henke (1998) also found that Texas

horned lizards selected recently burned areas compared

to areas with large litter accumulation because of

increased mobility. Pianka (1966) found that horned

lizards preferred open areas to sit and wait for their prey,

thus increasing foraging efficiency. Open habitats also

aid in thermoregulation by allowing horned lizards

exposure to direct solar radiation (Heath 1965). Finally,

horned lizards may select open habitats due to an

increase in food abundance, specifically of harvester ants.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 49

Smaller home ranges in the inactive season were

expected based on our observations of horned lizard

activity. Fair and Henke (1999) also found that home

ranges decreased in size as the summer progressed until

hibernation. Several reasons could explain seasonal

differences in activity and ranging behavior of horned

lizards. First, increased mobility of horned lizards

during late spring and early summer could be due to

mate-searching and nest-building activities. Horned

lizards on the CWMA typically emerge from hibernation

in early March or April and become highly mobile, often

moving > 100 m/day to reproduce, build nests, and lay

eggs. Second, as the summer progressed, the

temperatures rose to points that could be lethal to

horned lizards (Forrester et al. 1998); therefore, horned

lizard movements were likely constrained by

temperature in the inactive season. Third, the increase in

harvester ant abundance and activity in the inactive

season may have enabled lizards to move shorter

distances to find food, thereby reducing the ranging

behavior of the lizards.

Estimates on survival rates of horned lizards are

contentious due to the large number of censored lizards.

Munger (1986) found that Texas horned lizards in

southeastern Arizona had annual survival rates between

35.0 and 86.0 percent, whereas Fair and Henke (1999)

estimated 8-month survival rates (Mar-Oct) in southern

Texas to be lower (8.9 -54.0 percent). However, Pianka

and Parker (1975) suggested adult Texas horned lizards

have comparatively high survival rates. Our estimates

(25 - 62 percent) fall between those of Munger (1986)

and Fair and Henke (1999). Lower survival rates in the

heavily grazed sites suggested that heavier levels of

grazing increase the vulnerability of horned lizards to

mortality and may counteract increased prey abundance.

Burning did not affect summer survival rates of horned

lizards, though survival rates were higher in burned than

unburned sites during the inactive season (P = 0.03).

Higher survival rates in moderately grazed sites and

burned sites could be due to better juxtaposition of food

and cover, as supported by the home range and ant data.

The smaller home ranges, increased survival rates (at

least in the inactive season), and greater prey abundance

in burned pastures suggested a positive effect of fire on

the ecology of Texas horned lizards. The effect of grazing

was more complex. Survival was decreased in heavily

grazed pastures, but range size did not differ among

grazing level. Also, ant activity was generally higher in

the heavily grazed pastures, especially when coupled

with burning. Our comparisons to an ungrazed,

unburned control were limited by a small sample in the

control pasture. However, measures of lizard resources

(e.g., vegetation, ants) and performance (e.g., range size,

survival) were generally comparable between control

and treated pastures. An alternate-year burning regime

and stocking rates of livestock such as those

implemented by CWMA created suitable habitat for

Texas horned lizards in southern Texas.

Acknowledgments

Texas Parks and Wildlife Department and David

Synatszke deserve special recognition for providing the

financial support to undertake this project as well as for

providing excellent accommodations throughout the

field seasons. We thank Sigma Xi Grants-in-Aid of

research for partially funding this research. We express

our gratitude to both D. Rios and A. Gandaria for their

assistance with data collection in the field. A special

thanks to L. Burrow for her help in making hundreds of

custom made backpacks for horned lizards. We thank

Dr. Mark Payton for his statistical expertise and many

hours of time agonizing over my project. Our sincere

appreciation to J. Vessels for reading this manuscript.

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52 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Abstract.—Fire has long been a natural and

anthropogenic force shaping southern forests and their

fauna. Some species are attracted to recent burns. There

is little direct mortality of adult birds by fire, but

growing season fires may consume some nests. Fire

affects bird communities mainly through effects on

vegetation. Fires effective enough to limit understory

hardwood development would reduce habitat suitability

for associated birds, but would promote species

associated with grass-forb vegetation. In the long term,

fire would disfavor species associated with deciduous

foliage and favor species associated with pine canopies.

Introduction

For thousands of years fire has been a natural and

anthropogenic force shaping southern forests and their

fauna. Southern forests and the animals that inhabit

them evolved with fire, and fire has helped shape

southern systems. Lightning-set fires have been a

recurring force (Heyward 1939), natives in the region

used fire to manipulate vegetation and drive game for

harvest, and European settlers used fire for the same

purposes and to clear new land for their crops.

Pine savannahs of the southern Coastal Plain with their

grassy understories were fire adapted. Longleaf pine and

red-cockaded woodpecker are fire adapted species, and

fire played a role in perpetuating the upland oak forests.

Natural fires periodically still affect southern forests and

prescribed burning continues to be used for a variety of

purposes, including wildlife management. Currently,

prescribed fire is used extensively to maintain pine

ecosystems, and particularly to manage for the redcockaded woodpecker.

In this publication I treat the effects of fire on birds in

the South. There are some recent data on fire and birds

in the South. And inferences can be drawn from data

from other regions, general information about effects of

fire on vegetation, and bird-habitat relationships

(Dickson 1981).

Variability

Fires and their effects on systems are extremely variable.

Every fire and its effects are different; and usually there

is considerable variation within each fire. Fires and their

effects involve habitat conditions before burning and

suitability for a wide variety of different communities or

species; the intensity, periodicity, and seasonality of

fires; landscape context and unburned areas, and

numerous interactions. For example, a low intensity

winter fire with high relative humidity, high fuel

moisture, carried by a low wind over a moist forest floor

has little long-term impact. Conversely, intense

conflagrations have long term effects on systems and

their components. I approach this treatment by

describing how fire may affect forests, and how that may

affect bird species and communities.

Short Term Effects

Obviously, fire has the potential to kill animals; and

generally, animals with low mobility are the most

vulnerable. There is documentation of some mammals

perishing in fires (Bendell 1974), and probably some

reptile and amphibian mortality. But most animals can

escape fire by fleeing, finding unburned shelter, or

burrowing. There probably is little direct mortality of

birds from fire, although growing season fires probably

consume some nests located on or near the ground. For

example, Doerr (et al. 1970) observed that no ruffed

grouse (see Tables 1 and 2 for scientific names of

animals and plants, respectively) were directly killed by

a fire but grouse nests apparently were consumed. Direct

mortality of animals from fire on a broad population or

landscape scale is negligible.

Some birds actually are attracted to the heat and smoke

of fires, or to the burns shortly thereafter. A list of 77

species of North American birds that were attracted to

fire and smoke or observed on recent burns was

compiled by Komarek (1969). Wild turkeys and

mourning doves are attracted to new burns in search of

exposed insects and seed. Raptors, such as red-tailed

hawks, kestrels, other hawks, and owls have been

observed attracted to burns in search of prey (Stoddard

1963, Landers 1987). Other species attracted to recent

burns include purple martins, American robins, eastern

bluebirds, sparrows, pine warblers, common flickers,

and other woodpeckers.

Habitat

Generally fire consumes forest floor litter and sets back

plant succession; usually reducing smaller hardwoods in

favor of thicker-barked pines and herbaceous vegetation.

Small top-killed hardwoods usually resprout after

infrequent and/or cool season fires; but conversely,

small hardwoods can be severely reduced or eliminated

by repeated, and especially growing-season burns (Lewis

and Harshbarger 1976). Hardwood trees and shrubs

impede more light to the forest floor than pines;

therefore, burning severe enough to reduce hardwoods

would favor vegetation near the ground, especially

Fire and Bird Communities in the South

James G. Dickson1

1School of Forestry, Louisiana Tech University, Ruston, LA

71272
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 53

grasses and forbs. In upland oak-hickory stands burning

favors mid-successional oaks which resprout, over the

succession of more tolerant hardwoods.

Fire affects hardwoods and their suitability for birds in

other ways. Fire wounds on hardwood trees provide

entrance for decay, which over time may become

excavated into cavities by woodpeckers, and used by

primary and subsequently secondary cavity nesters.

Severe fire may kill trees and create snags that are used by

a variety of cavity-using wildlife. But conversely, standing

dead snags used by cavity nesters may be consumed by

fire.

Bird Communities

In the short term there is some site fidelity of birds;

many remain on the site in spite of burn-induced habitat

changes (Bendell 1974). In south Florida there was little

immediate effect of burning of a slash pine stand on the

bird community (Emlen 1970). Emlen concluded that

bird attachments to home ranges and familiar foraging

areas transcended species habitat selection immediately

following a fire.

Burning usually does affect bird communities through

habitat changes for several years postburn (e.g.,

Engstrom et al. 1984, White et al. 1999). Bendell (1974)

summarized a number of studies of effects of burning on

birds. He concluded that overall, the number of species

of birds increased after burns. The number of species that

fed on or near the ground increased the most, as might

be expected with the decrease of trees and the

proliferation of grasses, herbs, and shrubs after burning.

The number of tree canopy species decreased the most.

Some specific examples of the influence of fire on bird

communities illustrate. In a pine stand in northern

Florida that had been burned regularly, a number of

species dwindled over time with fire exclusion as dense

hardwood sapling stand developed (Engstrom et al.

1984). With 15 years of fire exclusion open habitat

species disappeared quickly, shrub associated species

increased then dwindled, and mesic forest species

invaded.

In a study of prescribed burned mature pine stands the

Georgia Piedmont avian species richness was similar

between burned and unburned stands, but about 3 times

as many species preferred the burned stands (White et al.

1999). Burned stands were much preferred over

unburned for nesting. But productivity was very low on

burned sites, with predation implicated as the major

cause of nest failure.

A somewhat similar pattern in the bird community was

observed with hardwood removal (WSI) and burning in

the Ouachita Mountains of Arkansas (Wilson et al.

1995). Species richness was similar among treatments.

However, bird densities were highest the second year

after WSI and burning, and lowest in unburned

controls. Densities of ground/shrub-foraging and

shrub-nesting species increased the most following WSI

and fire. Ground-nesting species were more abundant

in untreated stands.

Bird populations were tracked for 15 years after a

wildfire in a coniferous forest in California (Bock et al.

1978). After 8 years, the bird community was slightly

richer and more diverse in burned than unburned

forest. Open ground and brush species predominated

on the burned areas, whereas coniferous foliage feeders

predominated on the unburned area. Eight to 15 years

post burn ground foragers and cavity nesters declined

on the burned area. After 15 years the burned area had

become brushy and most fire-killed trees had fallen.

Bird density and diversity were similar on the burned

and unburned areas.

A wildfire in ponderosa pine forests in Arizona

drastically affected the bird community (Lowe et al.

1978). The number of birds increased the first year after

the burn, but decreased substantially from 5 to 20 years.

The abundance of timber gleaning, flycatching, aerial

flycatchers, tree foliage searching, ground and brush

foraging birds, and timber drilling birds changed over

time in response to the burn.

From 2 to 4 years after a Minnesota wildfire 54 species

decreased, 17 species increased, and 42 species

remained relatively unchanged (Niemi 1978).

Waterbirds, flycatchers, swallows, vireos, and warblers

decreased the most; while woodpeckers increased the

most.

Fire also affects birds other than during the breeding

season. For example, during winter in a northwestern

coniferous forest that was burned by a wildfire, the

number of bird species was similar between burned and

unburned stands, but composition was distinctly

different (Kreisel and Stein 1999). Trunk and bark

foraging species, mostly woodpecker, were twice as

abundant in burned stands than unburned, but

declined in abundance in subsequent years after the

burn.

Bird Species in Southern Forests

A few species, such as the red-cockaded woodpecker,

brown-headed nuthatch, and pine warbler are

associated closely with mature pine trees (Johnston and

Odum 1956). Fires severe enough to preclude

hardwoods and favor pines would favor these species.

Birds were contrasted in mature pine stands burned

regularly, in part, for the red-cockaded woodpecker,

versus unburned pine-hardwood stands the Georgia

Piedmont (White et al. 1999) and in the Ouachita
54 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Mountains of Arkansas (Wilson et al. 1995). In both

areas there were no red-cockaded woodpeckers in the

unburned stands with hardwoods, and significantly

fewer brown-headed nuthatches and pine warblers.

But most bird species in pine stands are associated with

non-pine vegetation. For example, in pine stands in

Georgia the hardwood understory largely determined

the nature of the bird community (Johnston and Odum

1956). And in eastern Texas middle-aged pine stands

with no hardwoods were virtually devoid of birds

(Dickson and Segelquist 1979). With fire, the

composition and status of bird communities would be

largely determined by habitat responses to burning

(Dickson et al. 1993).

Typically, frequent, intense, or growing season fire

reduces small hardwoods and shrubs, and precludes

sizable hardwoods from developing. The presence and

abundance of birds associated with hardwood canopy,

such as the yellow-billed cuckoo, great crested flycatcher,

tufted titmouse, red-eyed vireo, black-and-white warbler,

and worm-eating warbler would depend on the extent of

hardwood canopy, that could be affected by the fire

regime of an area. In the Ouachitas, black-and-white

warblers were more abundant in stands with hardwoods

than in stands where hardwoods had been controlled

(Wilson et al. 1995).

Several species, such as the Acadian flycatcher, Carolina

wren, hooded warbler, Kentucky warbler, and northern

cardinal are associated with shrub-level and hardwood

midstory vegetation (Dickson and Noble 1978). The

distribution and relative abundance of these species

could depend on the extent of appropriate habitat,

which could be determined by fire (Dickson 1981). As a

hardwood sapling subcanopy developed with fire

exclusion in a pine stand in northern Florida, species

associated with mesic woods i.e., yellow-cuckoo, wood

thrush, red-eyed vireo, and hooded warbler, increased

(Engstrom et al. 1984). In the Georgia Piedmont, the

northern cardinal was more abundant on unburned

sites, but the Acadian flycatcher and Carolina wren were

more abundant on burned sites (White et al. 1999). In

the Ouachitas, both the cardinal and Carolina wren

thrived in stands where hardwoods had been killed but

had not been burned (Wilson et al. 1995).

The reduction of hardwoods through fire would

promote the development of fire-maintained pine

savannahs with associated grass-forb and low shrub

vegetation, with abundant seeds and fruit (Lewis and

Harshbarger 1976). This situation would benefit species

associated with these conditions, such as the prairie

warbler, common yellowthroat, yellow-breasted chat,

field sparrow, Bachman’s sparrow, grasshopper sparrow,

eastern towhee, and indigo bunting (Johnston and

Odum 1956, Meyers and Johnson 1978, Dickson and

Segelquist 1979, White et al. 1999). This relationship is

supported by fire exclusion data from the Coastal Plain.

Stands developed from a pine savannah, to shrub, to

hardwood subcanopies (Engstrom et al. 1984).

Bachman’s sparrows were soon eliminated from the

stand. After a few years yellow-breasted chats, common

yellowthroats, and indigo buntings were virtually gone.

Eastern towhees and northern cardinals responded

positively to developing shrubs, but dwindled as

hardwood trees dominated. In the Ouachitas, chipping

sparrows, indigo buntings, and Bachman’s sparrows

were more common on WSI/burn plots, and prairie

warblers numbers increased with years post-burn as

hardwood shrubs developed (Wilson et al. 1995).

Grass-forb vegetation would also provide important

brood habitat for southern gallinaceous species:

northern bobwhites and wild turkeys. In the Ouachitas,

both bobwhites and turkeys were detected regularly on

the WSI / burn plots (Wilson et al. 1995).

As noted, there usually is much variation in fire effects

on vegetation due to differences in local factors such as

fuel loading, size, and moisture content; as well as local

topography, soils, and daily weather. Fires which result

in a variable habitat with some unburned patches would

probably enhance bird abundance and diversity, and

generally be positive for birds. For example, Roth (1976)

successfully predicted bird species diversity in

brushlands with a vegetation heterogeneity index. In the

South (Piedmont, White et al. 1999; Ouachitas, Wilson

et al. 1995), bird densities were higher in burned

treatments than in controls, probably due to the patchy

habitat resulting from fire. The eastern wood pewee, an

edge species, responded positively to fire in the

Ouachitas (Wilson et al. 1995). Other typical edge

species of southern forests which may benefit from

habitat variability resulting from fire would be: the great

crested flycatcher, and Carolina chickadee (Strelke and

Dickson 1980).

Fruiting

During fall and winter, fruits and seeds are a main diet

item of birds (Martin et al. 1951). Burning affects fruit

production and availability. Burning exposes seeds to

foraging birds in forest floor litter. For example, in the

Georgia Piedmont there were significantly more

herbaceous plant and legume seeds on prescribed

burned plots than on unburned plots the first growing

season after burning (Cushwa et al. 1966). And in the

Georgia Coastal Plain there was significantly less seed in

subsequent years after burning (Buckner and Landers

1979).

Short and long term effects of fire on fruiting are more

variable. In eastern Texas the first year after fire, the

number of plants with fruit was reduced by 72% (Lay

1956). Fruiting yaupon, holly, sweetleaf, and viburnum

were reduced. Fruiting dogwoods and American

beautyberry increased.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 55

After a couple of years post-fire, herbaceous vegetation is

gradually replaced by hardwood sprouts and shrubs.

Also, fruiting of shrubs recovering from fire and those

benefitting from reduction of vegetative competition

from fire increase. This fruit production benefits a

number of fruit consuming species, such as the wild

turkey and northern bobwhite (Landers 1987).

Arthropods

Also, although our knowledge of fire effects is limited,

arthropod populations and their interactions with

vertebrates certainly are affected by fire (Landers 1987).

Some species of insects are attracted to the flame and

smoke of fire, and reproduce in fire conditions (see Lyon

et al. 1978). Evans (1972) noted that Platypezid and

Empidid smoke flies, and some Cerambycid and

Buprestid beetles appear to be attracted to smoke; and

Melanophila oviposit in newly burned wood.

Effects of fire on arthropods on the forest floor seems

quite variable. It is generally conceded that invertebrate

soil fauna is reduced by fire; whereas surface insects are

less vulnerable to fire (Lyon et al. 1978). It has been

documented that parasites of wild turkey and northern

bobwhite are reduced in the short term by burning

(Dickson 1981).

Precautions

Red-cockaded woodpeckers peck cavity trees to produce

a resin flow around the cavity and down the tree bole.

Fire may ignite the resin up the tree bole of cavity trees

and may even burn out and gut the nest cavity(Conner

and Locke 1979). Therefore, surface fuel around cavity

trees may need to be raked away before burning.

Streamside zones (SZ) are strips of mature hardwoods

and pines along intermittent and permanent streams

traversing upland stands, often pine. Retention of

streamside zones (SZ) are important concessions for

wildlife in southern forests. These zones serve as travel

corridors for some species and provide important

mature habitat for birds (Dickson et al. 1995). These

moist hardwood-dominated sites usually will not carry a

fire. But with arid conditions and low fuel moisture fire

can a problem; and SZ should be protected from fire.

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Bock, C.E., M. Raphael, and J.H. Bock. 1978. Changing

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Conner, R.N., and B.A. Locke. 1979. Effects of a

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Dickson, J.G., and C.A. Segelquist. 1979. Breeding bird

populations in pine and pine-hardwood forests in

Texas. Journal of Wildlife Management. 43: 549-555.

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E. Franzreb. 1993. Effects of silviculture on

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Ortego. 1995. Streamside zones and breeding birds

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Doerr, P.D., Keith, L.B., and Rusch, D.H. 1970. Effects of

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The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 57

Table 1.—Scientific names of animals

Red-tailed hawk Buteo jamaicensis

American kestrel Falco sparverius

Wild turkey Meleagris gallopavo

Ruffed grouse Bonasa umbellus

Northern bobwhite Colinus virginianus

Mourning dove Zenaida macroura

Yellow-billed Cuckoo Coccyzus americanus

Common flicker Colaptes auratus

Red-cockaded woodpecker Picoides borealis

Great Crested flycatcher Myiarchus crinitus

Acadian flycatcher Empidonax virescens

Eastern wood-Pewee Contopus virens

Purple martin Progne subis

Carolina chickadee Parus carolinensis

Tufted titmouse Parus bicolor

Brown-headed nuthatch Sitta pusilla

Carolina wren Thryothorus ludovicianus

Wood thrush Hylocichla Mustelina

American robin Turdus migratorius

Eastern bluebird Sialia sialis

Red-eyed vireo Vireo olivaceus

Black-and-White warbler Mniotilta varia

Worm-eating warbler Helmitheros vermivorus

Pine warblers Dendroica pinus

Prairie warblers Dendroica discolor

Common yellowthroat Geothlypis trichas

Yellow-breasted chat Icteria virens

Kentucky warbler Oporornis formosus

Hooded warbler Wilsonia citrina

Northern cardinal Cardinalis cardinalis

Indigo bunting Passerina cyanea

Eastern towhee Pipilo erythrophthalmus

Grasshopper sparrow Ammodramus savannarum

Bachman’s sparrow Aimophila aestivalis

Chipping sparrow Spizella passerina

Field sparrow Spizella pusilla

Table 2.—Scientific names of plants

Longleaf pine Pinus palustris

Slash pine Pinus elliottii

Ponderosa pine Pinus ponderosa

Holly Ilex opaca

Yaupon Ilex vomitoria

Dogwood Cornus florida

Sweatleaf Symplocos tinctoria

American beautyberry Callicarpa americana
58 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Abstract.—Prescribed fire is being used extensively for

habitat management of non-game birds, although the

area burned today is small relative to the amount of

land that burned historically. Results of a non-scientific

questionnaire of public and private land managers in

the eastern U.S. revealed prescribed fire is being used to

provide winter, breeding season, and migration habitat

for at least 57 species of birds in 29 states. Increasingly

sophisticated application of fire will be necessary to

manage habitat for diverse bird species and other

organisms. Contemporary training courses on the use of

prescribed fire typically divide ecological effects of fire

on animals into two categories: direct effects alter the

animals’ physical condition and indirect effects are

mainly associated with changes in habitat. The

envirogram, a conceptual tool developed by

Andrewartha and Birch (1984) within their theory of the

environment, can be used to diagram multiple direct

and indirect effects of fire on bird species. Variables of

prescribed fire (i.e., ignition pattern, season, frequency,

etc.) can be matched within the envirogram to achieve

the desired management objectives for individual

species. We used a modified envirogram to examine how

prescribed fire is used in intensive management for the

northern bobwhite (Colinus virginianus) within the Red

Hills of north Florida and south Georgia. A coarsegrained approach to management involving multiple

species or an ecosystem may be more efficient and

sustainable than emphasis on a single species.

Introduction

Extensive use of fire to manipulate vegetation by Indians

disappeared from eastern North America centuries ago,

and natural fires (e.g., lightning-started) have been

reduced in the eastern United States by habitat

fragmentation, fire suppression, and alteration of

vegetation composition. Wildfires can be widespread

during times of prolonged drought—nearly one-half

million acres were burned in 2282 wildfires from 1 June

to 22 July 1998 in Florida (Lewis 1998)—but these

wildfires may behave very differently from natural fires

because of long periods of fire suppression. It is

probably not a coincidence that both the use of fire as a

management tool (Pyne 1982) and populations of bird

species that require habitat that is in early stages of

vegetation succession (Askins 1999) declined in the 20th

century. Current uses of prescribed fire can only be

considered a limited stabilization or reversal of that

trend.

Fire plays a critical role in ecosystem management as an

ecological process in many vegetation types in the

United States (Smith 2000). Fire has been long been

recognized as an essential disturbance in some

ecosystems, such as longleaf pine (Pinus palustris)

woodlands, which require a relatively high fire

frequency to suppress competing hardwoods and

maintain an open forest structure that fosters pine

regeneration. The endangered red-cockaded

woodpecker (Picoides borealis) depends on pine forest

structure and composition that are shaped by frequent

fire. In other ecosystems, such as oak woodlands,

savannahs, and barrens in the central hardwoods region

(Anderson and others 1999; Yaussy 2000), acceptance of

the role of fire continues to grow. The finer points of

prescribed fire will play an increasingly important role

in the success of natural resource management as

wildlands are reduced and multiple management

objectives increase.

Prescribed fire is pervasive in the growing emphasis of

modeling vegetation manipulations on natural

disturbances within the context of ecosystem

management (White and others 1999; Engstrom and

others 1999). According to a recent review, ecosystem

management is “...management driven by goals,

executed by policies, protocols, and practices, and made

adaptable by monitoring and research based on our best

understanding of the ecological interactions, and

processes necessary to sustain ecosystem composition,

structure, and function” (Christensen and others 1996).

In this definition, sustainability of resources is a

precondition of management for production.

Maintenance of viable populations of animals (e.g.,

game or endangered species) is one of many natural

resource goals that must be balanced within the

ecosystem management framework by a broader

“ecosystem composition, structure, and function.” From

this perspective, understanding the ecological relations

of a single species is a critical element of ecosystem

management within the constraints of ecosystem

sustainability.

In this paper we (1) summarize the results of a survey to

better understand the extent and nature of prescribed

fire applications to non-game bird management; (2)

examine the ecological basis for use of prescribed fire to

manage bird populations; and (3) examine application

of prescribed fire within the context of management for

a single-species, the northern bobwhite (Colinus

virginianus) within the Red Hills.

Burning for Birds: Concepts and Applications

R. Todd Engstrom1 and David J. Brownlie2

1

Tall Timbers Research Station, Tallahassee, FL 32312

2

U.S. Fish and Wildlife Service, Tallahassee, FL 32312
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 59

Use of Prescribed Fire for Bird

Management un the Eastern U.S.:

Survey Results

In the statement of purpose of this symposium, it was

stated “...with the exception of a few high-profile

threatened or endangered species, the use of prescribed

burning solely for nongame species or communities has

been a rare occurrence.” The dependence of some

threatened and endangered bird species (e.g., redcockaded woodpecker) on fire-maintained habitats is

well-established (James and others 1997). To obtain a

baseline evaluation of the current uses of prescribed fire

in relation to populations and communities of birds in

general, we surveyed 97 organizations that we thought

to be prescribed fire practitioners. We distributed our

survey electronically to contacts in the U.S. Fish and

Wildlife Service, USDA Forest Service, National Park

Service, state agencies (i.e., Forestry/Fire Protection,

Wildlife, and Natural Heritage/Endangered Species

programs), and The Nature Conservancy in 29 eastern

states. Fifty-eight of a total of 60 responses stated that

they burned property they manage, and, of these, 42

(72%) of the respondents stated that they burned

specifically to provide habitat for birds (Brownlie and

Engstrom 2001).

Fifty-seven bird species in 22 families (77% were

nongame species) were identified by respondents as

species that benefit from land management with

prescribed fire. (See Brownlie and Engstrom 2001 for a

complete list of species). Predictably, grassland birds,

such as the emberizid sparrows, were well-represented

and comprised 23% of the total; prescribed fire was also

used for management of gamebirds, such as quail, dove,

and ducks. Although our assessment of birds that are

managed with fire is general, it indicates that prescribed

fire is being used extensively for game and nongame

birds, not just threatened and endangered species and

that prescribed fire for management of avian habitat is

being applied in a wide variety of vegetation types

throughout the eastern United States. The diversity of

habitats that are being burned for a diversity of bird

species implies that many techniques of prescribed fire

are being used.

Concepts

Ecology provides the scientific foundation of ecosystem

management. One of the concepts critical to an

ecological understanding of the relationship between

birds and fire is plant succession. Christensen (1988)

developed a model of succession that involved four

phases: establishment, thinning, transition, and steady

state. In establishment, seedlings grow rapidly. In the

second phase, plants thin out during a period of intense

inter- and intra-specific competition. The third phase

involves gaps created in the canopy and initiation of

small-scale recruitment. In the last phase, the plant

community reaches a steady state in which

establishment, thinning, and gap formation occur in a

mosaic of patches. The scale and type of disturbances

influence the size and nature of plant response to the

patches. It is important to note that disturbance is

implicit in natural plant community development. The

stability of the “steady-state” is dependent on a variety

of factors. Alteration of the disturbance regime or plant

community composition (e.g., invasion of exotics) can

disrupt community equilibrium and potentially shift the

community into a new steady state (Wiens 1997).

Making theory relevant to managers is a substantial

challenge to ecologists and requires a constant search for

techniques that provide an interface between the two

disciplines: science and management (Rodgers 1997).

We use a simple conceptual technique, the envirogram

(Andrewartha and Birch 1984), as such an interface to

review some of the concepts that are essential for an

ecological understanding of the relationship between

fire and birds that is necessary for ecosystem

management.

Effects of fire on animals are frequently classified as

relatively short-term and direct (mortality or movement)

or long-term and indirect via plant succession and

alteration of vegetation structure (Smith 2000). This

simple dichotomy is expanded within the envirogram

(Figure 1; Andrewartha and Birch 1984). The

envirogram is composed of three parts: the focal

organism, the centrum (direct effects), and the web

(indirect effects). The ability of the focal organism to

survive and reproduce is influenced by four components

of the centrum—resources, mates, predators, and

malentities. The direct effects within the centrum are

affected by environmental influences within the web

that have increasingly less direct influence on the

organism of interest. Fire plays multiple positive and

negative roles within the centrum and the web for

different organisms. For birds, fire can be within the

centrum as a malentity, because it can kill an organism.

At the same time, fire can be used to improve the

structure of the habitat over a longer period.

In our envirogram, we match characteristics of

prescribed fire with aspects of the environment that

affect an animals’ ability to survive and reproduce

(Figure 1). Fire can be prescribed to increase or decrease

its effect as a malentity by manipulating method of

application and season. Season and severity can be

altered to influence availability of food and nesting

substrate. Area of burned acres on a landscape scale will

play an important role in creating or eliminating habitat

that will affect characteristics of the metapopulation of

the species of interest. Patchiness of a burn locally may

change the suitability of an area by altering shelter from

predators and proximate sources of food.

James and others (1997) used the envirogram as a

graphical technique to summarize the life history of the

red-cockaded woodpecker. In their envirogram, fire
60 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

played an indirect role in the amount of foraging habitat

(a resource) through the effects on the abundance and

distribution of hardwoods that compete with pines.

They also hypothesized that fire might play a role in

nutrient availability and population levels of cavity

competitors. Their research indicated that frequency and

season of fire were critical indirect influences on

measures of productivity of the woodpecker, although

they could not identify the exact causes of the

relationship. We used the envirogram for another bird

species that is managed intensively, the northern

bobwhite.

Birds and Prescribed Fire in

the Red Hills Landscape

The Red Hills physiographic region between Tallahassee,

Florida, and Thomasville, Georgia, includes

approximately 100 privately owned parcels in 310,000

acres of land that are managed for hunting (C. Ambrose

pers. comm.). White-tailed deer (Odocoileus virginianus),

mourning dove (Zenaida macroura), and ducks, primarily

wood duck (Aix sponsa) and ring-necked duck (Athya

collaris), are hunted, but northern bobwhite (Colinus

virginianus) is the raison d’être for the hunting estates.

Management for populations of bobwhite in the Red

Hills plays a notable chapter in the development of

prescribed fire as a management tool through the

contributions of Herbert L. Stoddard, Ed and Roy

Komarek, and Tall Timbers Research Station (Pyne 1982;

Johnson and Hale, this volume).

Approximately 90 to 100 bird species are resident, breed,

or overwinter throughout longleaf pine woodlands

(Engstrom 1993). Depending on ground cover

characteristics, approximately one third of the bird

species, including the northern bobwhite, may nest and

forage on or near the ground in longleaf pine

woodlands (Engstrom 1993). These ground-nesting and

ground-foraging species are presumably the species that

are most directly affected by fire. Many of the bird

species of the longleaf pine ecosystem are also common

in old-field pinelands, which are structural analogues of

longleaf pine woodlands (Engstrom et al. 1984),

although a detailed comparison has never been made.

Three species with affinities for open pine woodlands—

the northern bobwhite, Bachman’s sparrow (Aimophila

aestivalis), and red-cockaded woodpecker—disappeared

from an open oldfield pine woodland in northern

Florida (NB66, a 22-acre study area on Tall Timbers

Research Station) within 4 to 15 years after fire

exclusion (Figure 2). This plot is now a closed canopy

mixed hardwood-pine woodland with little groundcover vegetation.

Based on evidence from existing remnant uncultivated

tracts, pre-settlement upland vegetation on the hunting

estates was likely an open-structured woodland

dominated by longleaf pine with scattered species of

other southern pines and fire-resistant hardwoods (e.g.,

Quercus, Carya) with a species-rich understory (Platt et

al. 1988). Ground cover plant species richness in mesic

longleaf pine woodlands at a 1000 m2

 scale is among

the largest values reported in the temperate Western

Hemisphere (Peet and Allard 1993). Some parts of the

Red Hills, particularly in Florida, were converted to

agriculture in the first half, and turned fallow in the

latter half, of the 19th century (Paisley 1989). Agriculture

eliminated much of the longleaf pine and some of the

pyrogenic ground cover plants (e.g., wiregrass [Aristida

stricta]) on many of these old-field sites. Loblolly (P.

taeda) and shortleaf (P. echinata) pines frequently

replaced longleaf as overstory dominants.

Probability of 

species a1

to survive and 

reproduce

Resources

Predators

Mates

Malentities

CENTRUM WEB

2 1

(e.g., food, 

nesting 

substrate)

(e.g., shelter)

(e.g.,

metapopulation

dynamics, 

landscape ecology)

(e.g., direct mortality, 

rate of spread, severity)

[Fire characteristics--

intensity, patchiness, size 

of application, method of 

application, and season--

are tools within the web 

(indirect effects) at the 

disposal of managers to 

manipulate aspects of the 

centrum of species a1. 

Note: fire is part of the 

centrum as a malentity, 

because it can cause 

death.]

Figure 1.—Modification of a generalized

Andrewartha and Birch (1984) envirogram to

represent the various aspects of prescribed fire that

indirectly and directly affect the ability of a bird

species to survive and reproduce. Given the

characteristics of an avian population of

management interest prior to use of prescribed fire,

a manager can match ecological characteristics of a

bird species with the prescribed fire objectives.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 61

Typical historical fire behavior within longleaf pine

woodlands is believed to have been frequent (1 to 3 year

fire intervals) and low intensity that rarely killed mature

trees (Myers 1990). These fires reduced aboveground

herbaceous vegetation biomass to ash and killed or

topkilled most hardwood stems. Grasses and forbs

readily resprout after these low intensity fires. The

season of natural fires, based on lightning-started fires,

is strikingly different from the seasonal application of

prescribed fire for management of northern bobwhite

populations on the hunting estates (Figure 3).

Prescribed fires on the hunting estates are typically

initiated immediately after the end of quail hunting

season in late-February , March, or early April, whereas

lightning-started fires occurred most frequently in May

or June (Komarek 1968; Brennan and others 1995). The

choice of season for prescribed fire in the Red Hills is

based on the desire to avoid burning during the hunting

season, minimize loss of quail nests to fire, take

advantage of dry vegetation and relatively cooler, drier

weather, and prepare for timber management activities.

Many of the prescribed fires in the Red Hills are small to

intermediate scale (10 to 200 acres) although sometimes

as much as 1000 acres may be burned on a single land

ownership in a day. Landscape patterns of Red Hills

prescribed fire is typically controlled by using roads or

plowed lanes as firebreaks. Fires are mostly conducted

during the daytime (especially in Florida) under low

humidity, and light wind conditions, but nighttime fire

is permitted in Georgia. These occasional nighttime fires

often result in patchy low-severity burns. Flanking,

backing, and head fires are used depending on the

objectives of the managers. Ignition is typically with a

drip torch along roads from a truck or ATV (all-terrain

vehicle). Fire permits are required and are issued on a

daily basis by state (Florida and Georgia) forestry

departments.

Prescribed fire is used extensively to manage habitat to

produce high populations of northern bobwhite in the

Red Hills landscape in southern Georgia and northern

Florida. A modified envirogram reveals how fire directly

and indirectly influences northern bobwhite fitness

(Figure 4). This envirogram only includes factors that

are associated with fire. For example, the effects of fire

on mate availability are unclear and therefore left blank.

Resources in the centrum of the bobwhite envirogram

that are influenced by fire are mainly foods: seeds, leaves

and insects. Soil disturbance (harrowing) in food plots

and agricultural fields and fire are used to manage the

composition and structure of vegetation. Soil

disturbance creates suitable substrate for weedy plant

species that provide the preferred foods of quail. Woody

species tend to suppress grasses (Wilson 1998). Fire

frequently kills hardwood saplings or aboveground

stems that would radically alter the structure and

composition of the ground cover plant community,

particularly the grasses. Fire frequency, severity, and

season affect vegetation composition and age, which are

important for herbaceous insects that are food for young

bobwhite quail. Various hawks (e.g., Accipiter and Buteo)

are predators of bobwhite and the bobwhite requires

brush for escape cover. Patchiness of burn, although

difficult to obtain predictably, could provide essential

cover. The season for fire also might have a strong affect

on visibility of quail to hawk predators. Eliminating

cover by fire during hawk migration in the spring, when

numbers of hawks are at their peak, could increase

predation by hawks. On the other hand, delaying

burning until later in the spring to minimize predation

by migratory hawks creates the potential of increasing

the loss of bobwhite nests directly to fire. The relative

importance of loss of some adults to predation to loss of

nests to fire to bobwhite fitness should inform

management on the preferred season of fire.

Figure 2.—Changes in abundance of

northern bobwhite (dotted), redcockaded woodpecker (dashed), and

Bachman’s sparrow (solid) in an oldfield

pine woodland over a 15-year period in

response to fire exclusion (adapted from

Engstrom et al. 1984).
62 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Examination of this envirogram reveal some potential

difficulties in the long-term use of fire within the

framework of short-term maximization of bobwhite

numbers, particularly in the interaction between soil

disturbance, vegetation composition, and fire. Soil

disturbance—depending on its scale and intensity—may

cause the loss of native, pyrogenic grasses and forbs,

particularly wiregrass in the Red Hills. Creation of food

plots, small agricultural fields, and harrowing have the

simultaneous effects of generating preferred foods of the

bobwhite and eliminating the ground cover plant

species that are best adapted to carry fire. The grasses

and forbs that replace the pyrogenic species apparently

are less able to compete with invasive hardwoods. The

scale and severity of soil disturbance are likely to be

influential in ground cover plant species recovery.

The tradeoffs between intensive management for the

bobwhite and pyrogenic ground cover emerge in the

direct and indirect effects of the environment on

bobwhite fitness (Figure 4) and can also be seen within

the framework of coarse and fine filters (Agee 1999).

Management for short-term increases in bobwhite

numbers (fine filter) through soil disturbance alters the

ground cover composition that best supports prescribed

fire. Impairing the competitive balance between grasses

and woody plants through harrowing or plowing will

ultimately limit the effectiveness of prescribed fire as the

principal form of disturbance within the ecosystem. A

coarse-filter approach in which the natural ground cover

composition is left undisturbed (e.g., retention of

wiregrass) and, to a lesser extent, the season of fire is

shifted to the growing season within the Red Hills may

be necessary to prevent ecosystem instability and

potentially loss of ecosystem composition and function.

Conclusions

Use of prescribed fire has been challenged for reasons of

public health and safety. Challenges have also emerged

for ecological reasons, because fire has positive or

negative effects on different species (e.g., long-distance

migratory songbirds vs. red-cockaded woodpecker or

rare butterflies vs. grassland birds). These potential

sources of conflict will require increasingly sophisticated

use of prescribed fire to achieve multiple ecological

objectives. The tools at the disposal of the manager who

employs fire (severity, season, size, etc.) need to be

matched carefully with our best knowledge of the

environmental influences of one or more species of

interest.

• One current framework for understanding the

effects of fire on birds (see Smith 2000) follows the

theory of the environment developed by

Andrewartha and Birch (1983). The envirogram is a

useful graphical technique that captures the

multiple direct and indirect effects of fire—both

positive and negative—in the life history of an

organism.

• The envirogram is a single-species approach that

could be used in multi-species conservation

planning by selecting indicator species and

examining the location and nature of fire within

those envirograms. The complexity of managing for

multiple species suggests that a coarse-grained

approach may be necessary for long-term

sustainability.

• The longleaf pine ecosystem is largely a grassland

that has largely been eliminated and degraded. Fire

exclusion has dramatic effects on the avian

communities of longleaf pine woodlands, and

groundcover disturbance, habitat fragmentation,

and ecotone disruption have more subtle, long-term

effects on birds. Within relatively intact longleaf

pine forests, the frequency, season, ignition pattern,

and intensity of fire can be manipulated to obtain

the desired structure and function.

Figure 3.—Frequencies of fires started by

lightning (solid bars) and prescribed fires

initiated for northern bobwhite management

(open bars) in the Red Hills region of north

Florida and south Georgia (adapted from

Brennan and others 1995 with permission).
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 63

Acknowledgments

W. Wilson Baker and Robert Crawford collected the data

for the study of NB66.

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Survival and 

reproduction of the

Northern Bobwhite 

CENTRUM

WEB

3 2 1

Resources

Predators

Mates

Malentities

Seed and leaves, 

especially from weeds 

and plants along field 

margins

Insects for young and 

brooding females

Various hawks

Fire (season, ignition 

pattern, rate of 

spread, severity)

Vegetation 

composition

Fire (season, 

patchiness)

Fire (season, 

frequency, 

severity)

Soil 

disturbance

Escape cover, 

brush,

ecotones

Vegetation 

composition

Fire (season, 

frequency, 

severity)

Figure 4.—An abbreviated envirogram for the northern

bobwhite on hunting estates in the Red Hills region of

north Florida and south Georgia that includes all

occurrences of fire in the web and centrum.
64 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

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biology of grasses. Cambridge University Press,

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ecosystem management. R.C. Szaro, N.C. Johnson,

W.T. Sexton, and A.J. Malk, eds. Elsevier Science,

Oxford: 281-312.

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on fire, people, and the central hardwoods

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Tech. Rep. NE-274. Newtown Square, PA: U.S.

Department of Agriculture, Forest Service,

Northeastern Research Station. 129 p.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 65

Abstract.—The shelterwood-burn technique is a novel

method for regenerating oak-dominated stands on some

upland sites while simultaneously minimizing

undesirable hardwood intrusion with prescribed fire.

Management options available within an oakshelterwood burn regime will create variably structured

habitats that may potentially harbor avian communities

of mature forest and early successional species (canopy

retention); grove-woodland species (post-harvest

prescribed burn) or shrubland species (total harvest).

We suggest that the management options associated

with shelterwood-burn silviculture offer viable

alternatives for managing songbird and timber resources

where oak-dominated stands are the desired goal in

upland southeastern sites.

Introduction

Songbirds have been the focus of many conservation

efforts as declines in populations of many species,

especially Neotropical migrants, have been recorded in

the eastern United States (Askins et al. 1990). While

numerous investigators have reported the effects of

various silvicultural treatments on songbirds (e.g.

Conner and Adkisson 1975; Webb et. al. 1977; Evans

1978; Crawford et. al 1981) there is a dearth of

information addressing the effects (real or potential) of

prescribed fire on songbirds in hardwood systems of the

Southeast.

Oaks, Quercus spp., are one of the most important food

and cover resources for forest wildlife in the Southeast

(Martin et al. 1956). A large number of songbirds,

including many species of Neotropical migrants, occupy

oak forest types in southeastern North America (Hamel

et al. 1982). Additionally, oaks are a valuable economic

commodity producing high quality timber for a variety

of uses. Because of its value, oak regeneration is a

priority on many upland sites. Shelterwood silviculture

is widely used to regenerate oak stands on upland sites

(Sander et al. 1983). It is employed so that the partial

harvests will reduce the dense shade that suppresses vital

root development of existing oak regeneration (Loftis

1990; Sander 1971).

By retaining the canopy and maintaining partial shade,

the rapid growth of shade-tolerant species such as

yellow poplar (Liriodendron tulipifera), is inhibited. The

added benefit of litter and soil disturbance during

harvest operations prepares seed beds for acorns and

oak seedling establishment (Cook et al. 1998).

However, because of a paucity of oak seedlings and

sprouts in mature oak stands and/or the inability of

existing oak stock to out-compete other vegetation,

shelterwood cuts alone are often ineffective in

promoting oak regeneration (Smith 1993; Lorimer

1993). As a result expensive pretreatment measures such

as herbicide application (Loftis 1990; Lorimer et al.

1994), low-intensity prescribed fires (Barnes and Van

Lear 1998; McGill et al. 1999), tree shelters (Potter

1988) and nursery stock plantings (Bowersox 1993;

Gordon et al. 1995; Schlarbaum et al. 1997) must be

implemented 5-15 years before the initial harvest. Such

treatments are unattractive to natural resource managers

and private landowners with limited budgets.

A more efficient means of regenerating oak stands

(hereafter referred to as the oak shelterwood-burn

technique) was developed jointly in 1993 by research

conducted by the Virginia Department of Game and

Inland Fisheries (VDGIF) and Clemson University

Department of Forest Resources (Clemson, South

Carolina). By burning two oak-dominated shelterwood

stands after an initial harvest (Keyser et al. 1996), the

regeneration of yellow poplar, red maple (Acer rubrum),

and sweetgum (Liquidambar styraciflua) was reduced by

67-90% while oak reproduction was reduced by only

11%. Subsequent studies of fire effects in oak-dominated

shelterwood stands in the Virginia Piedmont (Brose and

Van Lear 1998a; Brose et al. 1999) and the Northeast

(Ward and Gluck 199) showed the same trends in fire

resistance of oak and demonstrated the critical role of

fire intensity coupled with growing-season burns for

creating a cohort of tree seedlings dominated by oaks.

Although the oak-shelterwood burn technique was

originally implemented to improve the viability of oak

regeneration and the production of hardwood timber in

uplands, the conservation of biodiversity is often a goal

of many forest management initiatives. Because the

shelterwood-burn method is a novel technique and

could be adopted by increasing numbers of landowners

and natural resource managers, it will be important to

understand how wildlife communities might respond.

Here, we examine how various management options

implemented in oak-shelterwood burn sites could

influence the composition of songbird communities.

The potential influences of the management options

described here are related to the vegetative structure and

composition that will result from the three oakManagement Options for Songbirds Using the Oak Shelterwood-Burn Technique

in Upland Forests of the Southeastern United States

J. Drew Lanham1, Patrick D. Keyser2, Patrick H. Brose3 and David H. Van Lear1

1

Department of Forest Resources, Clemson University,

Clemson, SC 29634

2

Westvaco Forest Resources, Rupert, WV 25984

3

Northeastern Research Station, USDA Forest Service, Irvine,

PA 16329
66 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

shelterwood burn options and from inferences drawn

from other bird habitat studies conducted in upland

oak-dominated stands where silvicultural treatments

have created habitat conditions similar to those expected

in oak-shelterwood burns.

The Shelterwood Burn Technique

The oak shelterwood-burn technique is a three-step

process. First, an initial shelterwood cut leaves 50-60

dominant oaks per ha (11 to 12 m2 of basal area/ha).

The remnant stand of oaks is comprised of the best

stock to provide a vigorous regeneration cohort. Next,

the stand is left undisturbed for 3 -5 years while the

regeneration layer develops. After 3-5 years, a hot (flame

length > 1.0 m) growing-season fire is applied to the

stand, resulting in an oak-dominated regeneration

cohort. Each of these steps and the added option of

complete stand harvest after the development of a

strong regeneration cohort will create three variably

structured forest habitats that may be used by a wide

variety of resident and migrant songbirds.

Options for Songbird Management

Option 1: Canopy Retention

Canopy retention treatments provide two-age stands

twice during the shelterwood burn cycle. First, during

the phase when shade-tolerant hardwoods such as

yellow poplar and red maple dominate the advanceregeneration pool and second, after a satisfactory cohort

of vigorous, advance oak regeneration is achieved when

a portion or all of the residual overstory trees may be

retained for at least half of the next rotation. Retention

of a partial overstory during either phase may provide

sufficient canopy habitat and vertical structure for some

species of mature forest birds (Dickson et al. 1995).

Relative to other even-aged silvicultural methods,

canopy retention treatments would be the least intensive

and probably most similar to an uneven-aged mature

forest.

Crawford et al. (1981) surmised that timber

management strategies altered bird communities in

relationship to the degree of stand disturbance. They

predicted that partial harvests would provide sufficient

canopy cover to buffer complete species turnover from

mature forest to early-successional species observed in

clearcut forests. They further stated that partial cuts

would return more quickly to site conditions conducive

to mature forest species than would even-aged

treatments. These findings have been corroborated by a

number of other studies that have shown that although

populations of some forest-interior songbirds may be

reduced relative to an undisturbed stand due to habitat

alteration, increased nest predation, and parasitism

(Webb et al. 1977; Nichols and Wood 1995), these

species are generally not entirely eliminated and

population recovery may occur rapidly as the new forest

matures (Conner and Adkisson 1975; Askins and

Philbrick 1987).

Dickson et al. (1995) support the idea that the retention

of a residual canopy ( <50%) for several years after an

initial harvest can provide habitats for some mature

forest birds that would not inhabit stands managed

using traditional even-aged management technique. In

West Virginia, Nichols and Wood (1995) found that

two-age stands contained a greater density, richness,

evenness, and overall diversity of breeding birds than

early-successional and mature stands. Total density for

all Neotropical migrants was also highest in the two-age

stands. Densities of forest-interior species were not

different between clearcut, mature, and two-age stands.

The two-age stands had densities of interior-edge species

equal to or greater than the other two treatments. These

patterns can be explained, in part, by the occurrence in

two-aged stands of species normally associated with (a)

forest interiors: veery (Catharus fuscescens), American

redstart (Setophaga ruticella), and scarlet tanager (Piranga

olivacea), and (b) early-successional habitats: chestnutsided warbler (Dendroica pensylvanica), indigo bunting

(Passerina cyanea), and eastern towhee (Pipilo

erythrophthalmus). The co-occurrence of mature forest

and early-successional species within the same areas

indicated that two-aged stands might provide both types

of habitats for these species.

Annard and Thompson (1997) reported higher species

richness for breeding birds in stands treated by

shelterwood cuts than in clearcuts, group selection,

single tree selection, or uncut stands in the Missouri

Ozarks. The number of species detected were higher in

shelterwoods than in uncut controls or uneven-aged

stands. As with Wood and Nichol’s study, these

differences were attributed to the presence of a mixture

of early-successional and mature forest bird species

including blue-winged warbler (Vermivora pinus), and

prairie warbler (Dendroica discolor). These species

occupied shelterwood stands along with birds more

commonly associated with mature stands such as redeyed vireo (Vireo olivaceous), worm-eating warbler

(Helmitheros vermivorous), and Acadian flycatcher

(Empidonax virescens).

Nesting success must be considered in conjunction with

measures of density and diversity of breeding birds.

Nichols and Wood (1995) did not find any differences

in nest success among treatments in West Virginia. Nest

parasitism by brown-headed cowbird (Molothrus ater)

was not a major factor in their study with only eight of

246 nests parasitized and no differences in the number

of cowbirds found among treatments. Annard and

Thompson (1997) and Welsh and Healy (1993) found

similar results in Missouri and New Hampshire,

respectively. One must remain aware, however, as

patterns of predation and parasitism may vary

depending on the landscape context. The impact of

cowbirds and predators in extensively forested systems
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 67

tends to be lower than those in agricultural and

suburban landscapes (Wilcove 1985).

Canopy disturbance has been shown to benefit some

forest-interior bird species that have declined in some

regions. Some bird species that use early-successional

gaps within mature forests may decline in areas where

disturbances do not produce the regenerating groundlayer and shrub vegetation they prefer (Franzreb and

Rosenberg 1997). Shelterwood harvesting increases light

levels and soil disturbances that stimulate the growth of

low vegetative cover, i.e herbs/forbs/shrubs. In West

Virginia, Nichols and Wood (1995) found that the

Kentucky warbler (Oporonis formosus), wood thrush

(Hylocichla mustelina), American redstart, and black-andwhite warbler (Mniotilta varia) were 2-3 times more

abundant in two-age stands than in uncut controls.

The retention of 11-12 m2

 of oak basal area/ha (50-60

dominant oaks/ha) in shelterwood stands also provides

reliable acorn sources (Healy 1997). This is important

because the acorns provide seed sources for

regeneration. Acorns are also one of the most important

wildlife food resources as they are consumed by more

than 200 wildlife species throughout North America.

Among these are many species of songbirds (Martin et.

al. 1951; Beck 1993). Based upon the floristic structure

of stands expected after canopy retention treatments,

Table 1 lists bird species that are likely to occur in the

diverse two-age structure of these areas.

Option 2: Shelterwood Prescribed Burning

The second option in the oak-shelterwood burn scheme

is the use of periodic prescribed fire in partially

harvested stands. Among the three options discussed

here, this method is likely to be intermediate in its

effects on the songbird community. Ultimately, the shift

in species composition will vary depending on the

vegetative structure that results from the season,

intensity, and frequency of the prescribed burns.

Dormant-season burns produce low-growing, sprouting

regeneration of shrubs and trees and stimulate the

production of soft mast (Stransky and Rose 1984). These

responses may provide forage, cover and arthropod prey

for many songbird species (Dickson 1981).

Repeat dormant-season burning increases the

abundance of oak regeneration. Oak regeneration is

limited by additional fires and then released at intervals

by withholding burning treatments, creating patchy

stands in different successional stages. Dickson (1981)

surmised that in southern pine and pine-hardwood

forests, a patchwork of different successional stages

within a stand (or across a landscape) could enhance

bird diversity and abundance. This patchwork would

obviously be dependent not only upon the frequency

and intensity of fires but also on the size, topography

and site capability of the area burned. In stands

managed with dormant season fires that will allow the

proliferation of hardwood shrubs and trees underneath

an open canopy , bird communities are likely to be

comprised primarily of shrub nesting (e.g.white-eyed

vireo, Vireo griseus) and midstory species (e.g.wood

thrush) along with species more characteristic of open

canopy forests such as the yellow-billed cuckoo,Coccyzus

americanus, and blue-gray gnatcatcher (Pilioptila

caerulea). More so than other burning treatments,

dormant-season fires in oak-shelterwoods are likely to

produce vegetative characteristics and therefore bird

communities more similar to two-age canopy retention

stands.

Annual or biennial prescribed burning during the

growing season would create open hardwood

woodlands and savannas by gradually eliminating much

hardwood shrub and tree regeneration while stimulating

production of ground-level herbaceous vegetation (Thor

and Nichols 1973). Oak woodland and savanna habitats

were described as common landscape features by early

explorers and settlers who observed the Native

Americans’ extensive use of fire (Pyne 1982; Buckner

1983; Van Lear and Waldrop 1989). Over time, however,

oak savannas and woodlands and some of the wildlife

species associated with them have become rare. The

restoration of hardwood savannas and open woodlands

would probably shift bird guilds from mature forestinterior species to canopy and midstory dwelling, open

woodland and grove species such as great-crested

flycatcher (Myiarchus crinitus), eastern wood-pewee

(Contupus virens), orchard oriole (Icterus spurius) and

summer tanager (Piranga rubra). Although growingseason fires might benefit some bird species, others

could be negatively impacted by burns initiated so late

that nesting and other breeding activities are disrupted.

Therefore, spring burning should be judiciously prescribed

as early as possible in the season so that direct impacts on

nesting or breeding birds are minimized.

Fire intensity (“hot versus cool”) also affects vegetative

structure and therefore avian community composition.

In a study conducted in Alabama pine-hardwood

Piedmont sites, Stribling and Barron (1995) found a

greater abundance and diversity of birds in burned

stands subjected to cool fires, with canopy, shrub and

cavity nesters being most abundant. Canopy, shrub and

bark feeding species were also more abundant in cool

burn sites than in untreated stands. These differences

were attributed to the increased heterogeneity of

vegetative structure (patchiness) of treated areas.

Stribling and Barron (1995) found ground-foraging and

ground-nesting songbirds to be more abundant in pinehardwood stands treated with a hot, early-spring fire

than in those treated with cooler early-spring fires. They

attributed this response to the removal of litter, which

they hypothesized provided better foraging and nesting

areas for birds in those guilds.

Some residual damage can occur from prescribed

burning. Residual overstory trees, especially thin-bark
68 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Table 1.—Neotropical migrant birds associated with oak-shelterwood burn options in upland southeastern forests.

Canopy Retention Shelterwood Burning Complete Harvest

yellow-billed cuckoo (Coccyzus americanus)

whip-poor-will (Caprimulgus vociferus) X

chuck-will’s widow (C. carolinensis) X

ruby-throated hummingbird (Archilocus colubris)X X

Acadian flycatcher (Empidonax virescens)

eastern kingbird

(Tyrannus tyrannus)

great-crested flycatcher (Myiarchus crinitus) X

eastern wood-pewee

(Contupus virens)

gray catbird (Dumatella carolinensis) X

wood thrush (Hylocichla mustelina)

veery (Catharus fuscescens)

blue-gray gnatcatcher (Pilioptila caerulea)

yellow-throated vireo (Vireo flavifrons) X

red-eyed vireo (V. olivaceous)

white-eyed vireo (V. griseus)

Blackburnian warbler (Dendroica fusca)

black-throated blue warbler (D. caerulescens)

black-throated green warbler (D. virens)

cerulean warbler (D. cerulea)

chestnut-sided warbler (D. pensylvanica) X

prairie warbler (D. discolor)

yellow-throated warbler (D. dominica) X

yellow warbler (D. petichia)

blue-winged warbler (Vermivora pinus)

golden-winged warbler (V. chrysoptera)

American redstart (Setophaga ruticilla)

black-and-white warbler (Mniotilta varia)

common yellowthroat (Geothlypis trichas)

hooded warbler (Wilsonia citrina)

Kentucky warbler(Oporonis formosus)

northern parula (Parula americana)

ovenbird (Seiurus aurocapillus)

Louisiana waterthrush (S. motacilla)

worm-eating warbler (Helmitheros vermivorus)

yellow-breasted chat (Icteria virens)

orchard oriole (Icterus spurius) X

Baltimore oriole (I. galbula) X

scarlet tanager (Piranga olivacea)

summer tanager (P. rubra) X

indigo bunting (Passerina cyanea) X

blue grosbeak (Guiraca caerulea)

Habitat associations inferred from “Primary Habitats” and “Key Habitat Requirements” designations by Hamel et al. (1982): Oakshelterwood analogs are as follows: canopy retention = sapling poletimber-sawtimber; shelterwood burning =grass-forb, sawtimber;

complete harvest =seedling-sapling.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 69

species such as maples (Acer spp.) and yellow poplars

and those with slash accumulations at their bases (Brose

and Van Lear 1999) are prone to fire-kill or damage.

However, the creation of dead and dying trees (snags)

provide important foraging sites for woodpeckers and

other bark gleaning species such as the black-and-white

warbler. Snags also provide perching/hawking sites and

roosting/nesting habitats. Larger sized snags are valuable

nesting habitats for both primary cavity excavators

(woodpeckers) and secondary cavity nesters including

Neotropical migrants such as the great-crested flycatcher

(Lanham and Guynn 1993). In addition to the valuable

functions snags, downed logs and other coarse woody

debris (e.g. tree tops, fallen limbs) provide for birds,

these features provide habitat for forest-floor-dwelling

arthropods, herpetofauna, and small mammals (Hanula

1996; Loeb 1996; Whiles and Grubaugh 1996) . These

provide food resources for songbirds and gamebird

species such as wild turkey, Meleagris gallopavo, and

northern bobwhite, Colinus virginianus. Coarse woody

debris also helps to prevent erosion in steep terrain by

slowing overland water flow (Van Lear and Danielovich

1987) and builds soil as it decays by slowly releasing

nutrients and organic matter. These actions will affect

the structure and composition of vegetation and

ultimately the avian community on a harvested site.

Because fire in forested stands can have such varied

effects, a wide variety of bird species may be supported

based upon fire frequency, intensity and various site

characteristics. Because most natural resource mangers

and private landowners will be primarily concerned

with the production of open, oak-dominated

woodlands, Table 1 lists some species likely to occur in

understory (growing-season) burned treatments that

result in park-like oak woodlands.

Option 3: Overstory Removal

A third option is harvesting all of the residual overstory

trees. This approach creates even-aged, early-successional

hardwood habitat and is the most intensive of the

options described here. In the initial year after harvest,

stands without an overstory will undergo a dramatic

turnover in species. Species such as indigo bunting and

field sparrow, Spizella pusilla, are common in

regenerating hardwood stands during these initial grassforb and seedling-sapling stages (Evans 1978). In

subsequent years tree saplings and shrubs increase the

vertical structure within a regenerating stand at which

point avian diversity and abundance levels can surpass

those found in mature stands (Conner and Adkisson

1975; Thompson and Fritzell 1990). Thompson et al.

(1993) attributed this peak in diversity and abundance

to increases in vegetative vertical structure and

horizontal patchiness within and among stands. In

southeastern uplands, regenerating seedling-sapling

hardwood habitats are preferred by shrub-scrub species

such as prairie warbler, yellow-breasted chat, Icteria

virens, and chestnut-sided warbler. Table 1 lists some

Neotropical migrant bird species typical of regenerating,

early-successional hardwood stands.

As regenerating stands age to form closed canopy

sapling-poletimber stands, species richness and

abundance frequently decreases to levels below younger

shrubland and older mature forest habitats (Conner and

Adkisson 1975). However, some forest-interior

songbirds such as black-and-white warbler and wood

thrush will begin using pole stands at this stage (Conner

and Adkisson 1975; Askins and Philbrick 1987).

Discussion and Conclusions

The hardwood forests of eastern North America are one

of the largest broad-leaved, deciduous ecosystems in the

world (Hicks 1997). Among these ecosystems, upland

oak-dominated types are among the most widespread

and important as economic and ecological resources.

The songbird communities dependent upon these

habitats in the Southeast include a large number of

Neotropical migrants (Hamel et al. 1982; Thompson

and Fritzell 1990). Since many of these species are

declining, the management of their habitats has become

a conservation priority. Although the prevailing

songbird conservation paradigm in many eastern

hardwood-dominated forests has been to limit harvests

to single/ group-tree selection or eliminate cutting

entirely, thousands of hectares of oak-dominated forests

occur on private lands where wildlife conservation goals

may be secondary to timber management priorities. This

means that no cut and selection cut options are often

unrealistic. Innovative management is needed to satisfy

both goals.

Burning as a silvicultural technique in southeastern

forests has traditionally been used in pine stands.

Conversely, it has been regarded as a disturbance to be

prevented in hardwood forest management. The

shelterwood burn technique has been shown to be an

effective method for regenerating oak-dominated stands

in the southeastern Piedmont (Van Lear and Brose

1999). A number of other studies conducted in two-age

shelterwood systems (e.g. Nichols and Wood 1995;

Annard and Thompson 1997) have proven that

shelterwood techniques offer a viable option for

songbird conservation/management. Additionally,

Stribling and Barron (1995) showed that fire could also

play a positive role in forest bird management. We

suggest that stands managed in different stages of the

shelterwood-burn process across a landscape would

offer habitats similar to those two-age and burned

stands. The result, we believe, will be a diversity of

habitats attractive to forest-interior, edge-interior, open

woodland and early-successional shrubland species.

 While wildlife and timber production goals are

frequently in opposition, the ability to reliably

reproduce oak-dominated stands using a less intensive
70 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

form of even-aged management like the oakshelterwood burn technique and the associated options

might prove to be a strategic tool for both wildlife

conservation and sustainable timber production in

southeastern uplands. We do not suggest that the oakshelterwood burn system offers solutions for conserving

every songbird species within a given stand. We do

suggest, however, that this technique offers novel

opportunities for sustainable timber production and

effective songbird conservation across southeastern

landscapes where both are management objectives.

Acknowledgments

This work was supported by the Virginia Department of

Game and Inland Fisheries and the Clemson University

Department of Forest Resources. We also thank Maria

Whitehead, Ph.D. Graduate Research Assistant for

editing this manuscript.

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Rep. SE-69: 45-50.

Ward, J.S.; Gluck, E. 1999. Using prescribed fire to

release oak seedlings from shrub competition in

southern Connecticut. In: Stringer, J., Loftis, D., eds.,

Proceedings 12th Central Hardwoods Conference.1-3

March 1999, Lexington, KY. USDA Forest Service Gen.

Tech. Rep. SRS-24: 283-287.

Webb, W.L.; Behrend, D.F.; Saisorn, B. 1977. Effect of

logging on songbird populations in a northern

hardwood forest. Wildlife Monographs. 55: 6-35.

Welsh, C.J.; Healy, W.M. 1993. Effects of even-aged

timber management on bird species diversity and

composition in northern hardwood forests of New

Hampshire. Wildlife Society Bulletin. 21: 143-154.

Whiles, M.R.; Grubaugh, J.W. 1996. Importance of

coarse woody debris to southern forest

herpetofauna. In: McMinn, J.W., Crossley, D.A., eds.,

Biodiversity and coarse woody debris in southern

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Forest Service Gen. Tech. Rep. SE-94: 94-100.

Wilcove, D. 1985.Nest predation in forest tracts and

the decline of migratory songbirds. Ecology. 66:

1211-1214.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 73

Abstract.—Shortleaf pine-bluestem (Pinus echinata Mill.-

Andropogon spp.) restoration for red-cockaded

woodpeckers (Picoides borealis) has been underway for

more than 2 decades on the Ouachita National Forest,

Arkansas. Restoration efforts consist of modifying stand

structure to basal areas similar to presettlement times

and reintroduction of fire. This is accomplished through

midstory and codominant tree removal (wildlife stand

improvement -WSI) and dormant season prescribed fires

on a 3-year cycle. Concern has been expressed about the

influence of this type of management on non-target

species, specifically small mammals and breeding birds.

Control stands (no thinning or fire) were characterized

by closed canopies and dense midstory with little

understory vegetation. WSI-treated stands were

characterized by open canopies, little midstory and an

increase in herbage production by 3-7 fold depending

on whether or not stands had been burned and time

since the stands had been burned. Woody cover after

WSI followed a predictable increase with additional

growing seasons since prescribed fire. Total community

abundance, species richness, and diversity of small

mammal and breeding bird communities were

dramatically increased by restoration. No small

mammal species were adversely affected by restoration

treatments. Pine-grassland obligate songbirds increased

following WSI and many of these showed significant

increases directly attributable to fire. Bird and small

mammal habitat associations are also described.

Introduction

Shortleaf pine (Pinus echinata Mill.) -grassland habitats

were once a prevalent landscape component in the

Ouachita Mountains (Nuttall 1980; Foti and Glenn

1991; Masters et al. 1995). Frequent fire maintained

these woodlands as distinctly open, pine-dominated

communities (Jansma and Jansma 1991; Masters et al.

1995). Fire suppression has led to the replacement of

pine-grassland woodlands with closed canopy pinehardwood forest types throughout the southeastern

United States (Waldrop et al. 1992). To improve habitat

conditions for the endangered red-cockaded

woodpecker (see list of common and scientific names

for bird species), the U. S. Forest Service began thinning

midstory and some codominant pine and hardwood

trees in 1979. This cultural practice, wildlife stand

improvement (WSI), is followed by dormant-season

prescribed fire every 3 years (U.S.D.A. 1985). In the

early 1990s, application of these practices was

broadened to an ecosystem approach for restoration of

the shortleaf pine- bluestem (Andropogon spp.)

community on the Ouachita National Forest.

For landscape and ecosystem restoration, quantitative

knowledge of historical vegetation patterns across the

landscape is essential in order to develop suitable

restoration targets. Historical land use documents such

as General Land Office (GLO) survey notes have

successfully been used to describe presettlement and

settlement landscapes (Cottam 1949, Lorimer 1980, Foti

and Glenn 1991). Analysis of GLO data in the Ouachita

Highlands of Arkansas (Foti and Glenn 1991) and later

in Oklahoma (Kreiter 1995) provided targets for stem

density, basal area and information on tree species

composition for renewal of the pine-bluestem

community.

Birds and small mammals have often been used as

representative taxa to determine vertebrate wildlife

responses to forest management practices (Morrison

1992, Masters et al. 1998). Bird distribution and

abundance is largely determined by vegetation structure

(MacArthur and MacArthur 1961; Wiens 1974; Roth

1976) and plant species composition (Smith 1977; Rice

et al. 1983; Brennan and Morrison 1991). As well,

habitat suitability for small mammals may be altered by

forest management practices that modify primary

production or vegetation structure (Masters et al. 1998).

The relationship between habitat attributes and bird

species richness and abundance has been demonstrated

in a number of forest types (e.g., Karr and Roth 1971;

Willson 1974; James and Wamer 1982; Yahner 1986)

and also for many small mammals (e.g., Dueser and

Shugart 1978; Grant and French 1980; Root et al. 1990;

Tappe et al. 1994). While several studies have examined

bird and small mammal-habitat relations following fire,

none we are aware of have sought to determine

population response and habitat characteristics

important to bird or small mammal species in restored

fire-dependent pine-grassland communities other than

Wilson et al. (1995) and Masters et al. (1998).

Pine-grassland restoration alters plant community

structure and composition that ultimately affect the

breeding bird and small mammal communities. Public

concern has arisen over the impacts of management for

Influence of Ecosystem Restoration for Red-cockaded Woodpeckers

on Breeding Bird and Small Mammal Communities

Ronald E. Masters1, Christopher W. Wilson2, Douglas S. Cram1, George A. Bukenhofer3 and Robert L. Lochmiller4

1

Department of Forestry, Oklahoma State University,

Stillwater, OK

2

The Nature Conservancy, Tulsa Oklahoma 74114

3

USDA Forest Service, Atlanta, GA 30309

4

Deceased
74 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

a single species on other members of the vertebrate

community. Detailed information on population

response and habitat relations in fire-dependent

woodlands is necessary to accurately assess the impacts

of WSI and prescribed fire in woodland restoration and

to determine relative success of efforts. Because

restoration efforts have been underway for a relatively

short period of time, the plant community may be in a

state of transition as additional burning cycles are

accomplished and as the extent of restored stands

increases. Therefore we followed up on our initial efforts

(Wilson et al. 1995) to assess the breeding bird

community. Our objectives were to document relative

abundance in untreated and treated stands and to

determine plant community structural and

compositional attributes that best explained breeding

bird and small mammal abundance.

Materials and Methods

Study Area

Our study stands were within a >60,000-ha area

proposed for pine-bluestem restoration in the westcentral Ouachita Mountains in the Poteau, Mena, and

Cold Springs Ranger districts of the Ouachita National

Forest, Scott County, Arkansas. Topography was

mountainous with numerous east-west trending ridges

and valleys. We initially focused on pine-bluestem

restoration stands with active woodpecker clusters or

evidence of past woodpecker occupation in the vicinity

(Neal and Montague 1991), but restoration efforts have

now expanded outside of previously known cluster sites.

All stands consisted of second-growth forest with a

history of timber management and ranged in size from

14-61 ha. The study area was previously described by

Wilson et al. (1995) and Masters et al. (1996).

Experimental Design

We used a completely randomized experimental design

with 5 treatments replicated >3 times/year over 4 years

for breeding birds and 2 years for small mammals.

Treatments were control (n =14); WSI-no burn (WSI; n

= 14); WSI, first growing season postburn (WSIB1; n =

17); WSI, second growing season postburn (WSIB2; n =

14); and WSI, third growing season postburn (WSIB3; n

= 14). In 1992, we randomly chose 12 stands from a list

of treated stands (n = 61), in addition to 3 randomly

chosen control stands. In 1993, all WSI and WSIB3

stands were burned in early spring and therefore,

became WSIB1 stands; WSIB1 became WSIB2, and

WSIB2 became WSIB3. Controls remained the same,

and we added 3 WSI stands for 18 stands sampled in

1993. In 1999 and in 2000, we randomly chose 20

stands each year (4 replications per treatment). The total

number of stands for breeding bird work was 73 and 30

for small mammals. Details of experimental design for

small mammals are found in Masters et al. (1998).

Bird and Small Mammal Surveys

We quantified relative bird abundance in 1992 and

1993, using 6, 40-m-radius point count plots 130-150

m apart with >90 m between plot centers and stand

boundaries within each stand (Hutto et al. 1986).

Methods, abundance, and frequency data were reported

in Wilson et al. (1995). In 1999 and 2000, we followed

the same methodology except that we sampled 3-point

count plots in each stand. Analysis of previous years

data demonstrated that 3 bird plots adequately

characterized birds of greatest interest and the most

abundant birds in the community.

We quantified small mammals using live traps and

mark-recapture techniques in winter 1992 and 1993 (27

Dec - 4 Jan). We located 80 Sherman live-traps at 15 m

intervals on randomly located parallel transect lines

within each stand. Methods, abundance and trap-nights

of effort were reported by Masters et al. (1998).

Vegetation Composition and Structure

We measured woody species composition and forest

structure in 4 circular, 5-m radius subplots within each

bird point count plot from late May to early June, 1992

and 1993 to examine habitat relationships. The first

subplot was located at bird point count plot center. The

second subplot was placed in a random direction, 30 m

from point count plot center. The remaining 2 subplots

were placed 120o

 and 240o from the second, and 30 m

from point count plot center.

We determined shrub density by counting all woody

stems 0.5-1.4 m in height within each subplot. We

categorized stems >1.4 m in height into 8 diameter

classes. We recorded all trees, shrubs, and vines within

each subplot by species. We used a 10-factor prism to

estimate conifer and hardwood BA in all years (Avery

1967) at the center of each subplot. We determined

coniferous and deciduous canopy heights by measuring

heights of 5 random overstory trees in each category

with a clinometer. We estimated percent canopy cover

in all years using a spherical densiometer (Lemmon

1957).

We characterized the vertical vegetation profile in 1992

and 1993 with a 10-m pole divided into 1-m segments

at 9 points within each subplot. Points lay 2.5 m apart

along perpendicular lines that intersected subplot center.

We extended the pole vertically through the midstory at

each point. We recorded presence or absence of

deciduous, coniferous, and dead vegetation touching the

pole for each 1-m segment.

We measured herbaceous and woody biomass by the

harvest method (Cook and Stubbendieck 1986) over a

2-week period from late July to early August using 30 1-

m2

 plots at 30-m intervals along randomized transects in

each stand (after Masters 1991). To avoid bias caused by
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 75

influence from adjacent stands, we did not sample

within 50 m of stand boundaries. We clipped current

year’s growth of all vegetation to <2.5 cm height and

hand separated into grass, sedge, panicum, non-legume

forb, legume, and woody categories. We dried samples to

constant weight at 70o

 C in a forced-air oven before

weighing. Standing crop data were reported by Masters

et al. (1996). We grouped data into grasslike, forb, and

woody categories for bird and small mammal habitat

analysis.

Data Analyses

We used the Kruskal-Wallis nonparametric test to detect

treatment differences (P < 0.05) in stem densities by

diameter class, basal areas, canopy heights, percent

canopy cover, understory standing crop, and diversity

measures. We analyzed data across years with stand

(year x treatment) type III mean square as the error

term, and used the least significant difference (LSD) test

to separate mean ranks (P < 0.05)(Conover and Iman

1981). To further differentiate treatment effects on

breeding bird and small mammal indices of relative

abundance, we used specific orthogonal contrasts to test

for differences between control, WSI, and burned and

unburned stands. To better determine effects of pinegrassland restoration on breeding bird community, we

grouped species into 3 designations of habitat

occurrence: forest interior, forest edge (including some

open forest species), and pine-grassland (see Wilson et

al. 1995). For community level analysis we calculated

Morisita’s Index of Similarity (after Krebs 1989) to

determine comparative community similarity between

treatments. Because unequal sampling effort can bias

Morisita’s Similarity indices particularly with small

sample sizes (Krebs 1989), we randomly selected 3

point count plots to discard for 1992 and 1993 to

maintain equal sampling effort among all years in

sampled stands.

We performed canonical correspondence analysis (CCA;

ter Braak 1986, 1987) with the program CANOCO (ter

Braak 1988) to determine the influence of 16

physiognomy and 21 floristic measures derived from

vegetation samples on abundances of 27 bird species

(with > 1 individual/40 ha, Wilson 1994). We used

Pearson product-moment correlations to examine the

relationships of these same physiognomy and floristic

variables with small mammal population and

community parameters because CCA results were

inconclusive (Masters et al. 1998).

Results

Stand Characteristics

Midstory removal (WSI) reduced percent canopy cover,

conifer BA, and total BA (Table 1). Hardwood BA was

lower in burned stands than in controls. Pine canopy

height and hardwood canopy height were similar among

controls and treatments. Shrub diversity (stems <8 cm

dbh) was greatest in WSIB2 and WSIB3 stands. Tree

diversity (stems >8 cm dbh) was similar among

treatments (Table 1). Stand characteristics in 1999 and

2000 were not different than for 1992 and 1993.

Table 1.—Stand characteristics after Wildlife Stand Improvement (WSI) and growing seasons since burned on

Ouachita National Forest, Arkansas, May 1992 and 19931

 Treatment2

CONTROL WSI WSIB1 WSIB2 WSIB3

Parameter Mean(SE) Mean(SE) Mean(SE) Mean(SE) Mean(SE) P>F

Canopy cover (%) 84.1(1.3) a 65.1(2.8) b 67.9(1.9) b 70.4(1.9) b 69.3(2.4) b <0.001

Conifer BA (m2

/ha) 19.5(0.4) a 13.5(1.7) c 14.8(1.1) bc 17.3(0.7) ab 16.8(0.9) abc 0.015

Hardwood BA (m2

/ha) 6.9(0.3) a 4.4(0.9) ab 3.9(0.9) b 2.6(0.9) b 2.9(1.1) b 0.042

Total BA (m2/ha) 26.4(0.4) a 17.9(1.1) b 18.7(0.9) b 20.0(0.8) b 19.7(1.3) b <0.001

Pine canopy ht. (m) 19.5(1.0) 18.2(0.9) 20.2(0.7) 20.3(0.9) 19.3(0.9) 0.559

Hdwd canopy ht. (m) 13.7(0.7) 14.8(1.0) 14.7(1.0) 11.5(2.0) 12.2(1.0) 0.308

Tree diversity3 1.4(0.1) 1.1(0.2) 0.8(0.1) 0.7(0.1) 0.6(0.2) 0.072

Tree spp. Richness 9.2(1.4) 6.2(1.3) 5.6(1.0) 5.7(0.8) 5.0(0.9) 0.332

Shrub diversity3 2.6(0.1) b 2.6(0.1) b 2.4(0.1) b 2.8(0.1) a 2.9(0.1) a 0.001

Shrub spp. Richness 33.2(2.9) ab 36.5(3.7) a 24.9(2.7) b 34.5(1.5) ab 37.7(1.8) a 0.020

1

Row means followed by the same letter or without letters were not significantly different at the 0.05 level (LSD).

2

Control = no treatment; WSI = wildlife stand improvement, no burn; WSIB1 = wildlife stand improvement, first growing

season following burn; WSIB2 = wildlife stand improvement, second growing season following burn; WSIB3 = wildlife stand

improvement, third growing season following burn.

3

Shannon-Weaver Diversity Index.
76 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Density of woody stems <1.4 m increased in WSI stands

as a result of sprouting (Table 2). Stem densities in all

categories from 1.1-23.0 cm dbh decreased (P < 0.05)

following WSI. The application of fire following WSI

reduced stem densities for all diameter classes <8 cm

dbh (P < 0.05). However, stem densities 0.5-1.4 m in

height increased in WSIB2 and WSIB3 compared to

controls (Table 2). Fire did not affect densities of stems

>8 cm dbh.

In the vertical profile, WSI reduced total vegetation in all

levels 1-10 m (Figure 1). Wildlife stand improvement

reduced occurrence of conifers up to 6 m and reduced

hardwood occurrence in the 1-9 m strata (P < 0.05).

Dead vegetation increased from 0-1 m in all treated

stands. Fire reduced occurrence of hardwoods and slash

from 0-1 m and conifers from 1-3 m. By the third

season postburn, hardwood vegetation from 0-3 m

increased, largely from resprouting oaks and hickories

(Figure 1).

Wildlife stand improvement increased standing crop of

grasslikes, forbs, and woody vegetation <1.4 m by 3-7

fold depending on whether the stand had been burned

and the time since burned (Masters et al. 1996). Woody

shrubs initially decreased following fire, but became the

dominant component of the understory by the third

season postburn (Masters et al. 1996).

Breeding Bird Response

We observed a total of 68 species of breeding birds in all

stands over the course of our study. Community

composition was different between WSI treated stands

and control stands in all years except 1992 (Figure 2 and

3). For all years combined, burned stands were > 97

percent similar in bird species composition.

Comparison of the bird community in controls versus

that in individual treatments were < 84 percent similar

for all years combined. WSIB2 and WSIB3 stands had

the most similar bird communities consistently in

individual years and in all years combined (Figure 2).

Forest Interior

The black and white warbler, ovenbird and scarlet

tanager declined in density (P < 0.05) in treated stands

from thinning. Only the former 2 species had declines

directly attributable to fire (P < 0.05) (Table 3).

Although a tendency for decline in density was observed

for the whip-poor-will, its frequency of occurrence

within treated stands was significantly (P < 0.05) less

than in untreated controls.

Forest Edge

American crow was the only species in this group to

decline (P = 0.033) from WSI (Table 3). We found

Table 2.—Average woody stem density (stems/ha) after WSI and fire on Ouachita National Forest, Arkansas,

summer 1992 and 19931

Treatment

CONTROL WSI WSIB1 WSIB2 WSIB3

Height class,

diameter class Mean(SE) Mean(SE) Mean(SE) Mean(SE) Mean(SE) P > F

0.5-1.4 m 2,155(432)b 10,638(2,996)a 4,647(1,279)b 11,157(826)a 2,070(1,966)a <0.001

>1.4 m

<1 cm 1,002(255) ab 784(258) b 94(27) c 478(81) b 1,550(279) a <0.001

1.1-2.5 cm 218(40) a 46(11) b 14(8) c 16(6 ) c 112(50) ab <0.001

2.6-8.0 cm 703(57) a 71(17) b 26(5) c 63(30) bc 92(34) bc <0.001

8.1-15 cm 351(24) a 30(12) b 20(5) b 31(8) b 41(23 b <0.001

15.1-23 cm 114(9) a 42(9) c 70(16) bc 96(19) ab 61(11) bc 0.005

23.1-30 cm 115(20) 71(6) 86(16) 95(16) 87(21) 0.548

30.1-38 cm 107(5) 88(17) 93(14) 99(12) 106(19) 0.881

>38 cm 39(4) 43(7) 31(7) 33(7) 34(6) 0.763

Total >1.4 m 2,648(231) a 1,175(277) b 434(22) c 910(69) b 2,082(303) a <0.001

1

Row means followed by the same letter or without letters were not significantly different at the 0.05 level (LSD).

2Control = no treatment; WSI = midstory removal within the past 2 years and no burning of downed trees; WSIB1 = midstory

removal, late winter prescribed burn, in first growing season; WSIB2 = midstory removal, late winter prescribed burn, in

second growing season; WSIB3 = midstory removal, late winter prescribed burn, in third growing season.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 77

9-10

8-9

7-8

6-7

5-6

4-5

3-4

2-3

1-2

0-1

-10 -5 0 5 10

Height (m)

WSI-NB

9-10

8-9

7-8

6-7

5-6

4-5

3-4

2-3

1-2

0-1

-10 -5 0 5 10

Height (m)

WSI-B1

9-10

8-9

7-8

6-7

5-6

4-5

3-4

2-3

1-2

0-1

-10 -5 0 5 10

Height (m)

WSI-B2

9-10

8-9

7-8

6-7

5-6

4-5

3-4

2-3

1-2

0-1

-10 -5 0 5 10

Height (m)

WSI-B3

9-10

8-9

7-8

6-7

5-6

4-5

3-4

2-3

1-2

0-1

-10 -5 0 5 10

Height (m)

CONTROL

Hdwd Conif Dead

LEGEND

Figure 1.—Vertical vegetation profile measured within bird census plots of wildlife

stand improvement (WSI)-treated and untreated stands on the Ouachita National

Forest, Arkansas, May 1992-1993. Control = no treatment; WSI = midstory removal

(thinning) and without burning of logging slash; WSIB1 = midstory removal, latewinter prescribed burn, following the first growing season after fire; WSIB2 = midstory

removal, late-winter prescribed burn, following the second growing season after fire;

WSIB3 = midstory removal, late-winter prescribed burn, following the third growing

season after fire;
78 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

American goldfinch, blue-gray gnatcatcher, brownheaded cowbird, Carolina wren, white-breasted

nuthatch and yellow-breasted chat density increased (P

< 0.05) in response to WSI (Table 3). Many of these

species including the common yellowthroat (P = 0.037)

responded favorably to the addition of fire in WSI

treated stands with the exception of the Carolina wren.

This species increased with thinning (P = 0.014) then

declined following the addition of fire (P = 0.035).

Pine-grassland

All pine-grassland species increased (P < 0.05) in density

following either WSI or, WSI and fire except Bachman’s

sparrow and northern bobwhite (Table 3). However,

frequency of occurrence within treated stands for these 2

species was significantly higher (P < 0.05) than in

control stands.

Small Mammal Response

We observed a total of 9 small mammal species in all

stands over the course of our study. Community

composition was different between WSI treated stands

and control stands in all years (Masters et al 1998).

Total community richness, diversity and abundance were

lowest in untreated stands. White footed-mouse (see list

for scientific names) was the most dominant species and

short-tailed shrew and harvest mouse were the next

most abundant species (Masters et al. 1998). The whitefooted mouse and short-tailed shrew occurred in all

stands (Figure 4, Masters et al. 1998). Also occurring but

less frequently were golden mouse, eastern woodrat,

cotton mouse, hispid cotton rat, pine vole, and deer

mouse listed in order of total captures (Masters et al.

1998). The WSI and WSIB1 treatments had the greatest

number of species (Figure 4). We found that no species

declined in response to either thinning or fire (Masters

et al. 1998).

Bird-habitat Relationships

When CCA was constrained to 16 explanatory

physiognomy variables, eigenvalues for axes 1 and 2

explained 48.7% of cumulative variance of birdphysiognomy relations (P < 0.01) (Wilson 1994). We

identified 7 variables (P < 0.05) that explained 46% of

variation in breeding bird abundances. They were:

relative presence of midstory foliage (2-10 m) (16%),

relative presence of foliage 0-2 m (8%), density of

hardwoods > 23.1 cm dbh (5%), percent canopy cover

(5%), density of all woody stems >1.4 m in height

(4%), density of all woody stems < 1.4 m in height

(4%), and density of conifer trees > 23.1 cm dbh (4%).

When ordination was constrained to 21 floristic

variables, eigenvalues for axes 1 and 2 explained 40.1%

of cumulative variance in bird species data (Wilson

1994). We identified 4 variables that explained (P <

0.05) 35% of variation in breeding bird abundances.

100 9 0 8 0 7 0 6 0

Percent similar

WSIB1

WSIB3

WSIB2

CONTROL

WSI

1992

100 9 0 8 0 7 0 6 0

Percent similar

WSIB2

WSIB3

WSI

WSIB1

CONTROL

1993

100 9 0 8 0 7 0 6 0

Percent similar

WSIB2

WSI

WSIB3

WSIB1

CONTROL

1999

100 9 0 8 0 7 0 6 0

Percent similar

WSIB1

WSIB2

WSIB3

WSI 

CONTROL

2000

100 9 0 8 0 7 0 6 0

Percent similar

WSIB2

WSIB3

WSIB1

WSI

CONTROL

All Years

Figure 2.—Bird community similarity (Morisita’s Index

of Similarity) among treated versus untreated stands on

the Ouachita National Forest, Arkansas, May 1992-1993

and 1999-2000. Control = no treatment; WSI =

midstory removal (thinning) and without burning of

logging slash; WSIB1 = midstory removal, late-winter

prescribed burn, following the first growing season after

fire; WSIB2 = midstory removal, late-winter prescribed

burn, following the second growing season after fire;

WSIB3 = midstory removal, late-winter prescribed burn,

following the third growing season after fire.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 79

Figure 3.—Bird community model of the response of

important bird species illustrating succession windows of

suitable habitat among restored versus untreated stands on

the Ouachita National Forest, Arkansas, May 1992-1993

and 1999-2000. Control = no treatment; WSI = midstory

removal (thinning) and without burning of logging slash;

1st growing season = midstory removal, late-winter

prescribed burn, following the first growing season after

fire (WSIB1); 2nd growing season= midstory removal, latewinter prescribed burn, following the second growing

season after fire(WSIB2); 3rd growing season= midstory

removal, late-winter prescribed burn, following the third

growing season after fire (WSIB3).

Figure 4.—Small mammal community model of the

response of most species and general habitat relationships,

illustrating succession windows of suitable habitat among

restored versus untreated stands on the Ouachita National

Forest, Arkansas, May 1992-1993. Control = no treatment;

WSI = midstory removal (thinning) and without burning

of logging slash; 1st growing season = midstory removal,

late-winter prescribed burn, following the first growing

season after fire (WSIB1); 2nd growing season= midstory

removal, late-winter prescribed burn, following the second

growing season after fire(WSIB2); 3rd growing season=

midstory removal, late-winter prescribed burn, following

the third growing season after fire (WSIB3).

They were: Shannon diversity index of woody stems > 8

cm dbh (16%), number of oak (Quercus spp.) stems > 8

cm dbh (9%), current annual growth of woody biomass

< 1.4 m in height (5%), and the number of pine trees >

8 cm dbh (5%).

In the physiognomy CCA, variables most correlated with

axis 1 were percent canopy cover, density of hardwoods

> 23.1 cm dbh, midstory foliage, and density of woody

stems >1.4 m in height (Wilson 1994). This axis

separated species dependent on closed canopy and

dense midstory forest from species requiring open

canopy and fewer trees. Variables most correlated with

axis 2 were number of pine trees > 8 cm dbh, density of

all woody stems < 1.4 m in height, and relative presence

of foliage 0-2 m. In addition to species demonstrating

affinities for numbers of large conifers, this axis

separated species relating positively to understory

hardwood structure from species preferring little or no

understory hardwood structure (Wilson 1994).

In the floristics CCA, large oaks and tree diversity were

most correlated with axis 1. This axis separated species

with affinities for density and diversity of hardwood

trees from species preferring fewer oaks and lower tree

diversity. Variables most correlated with axis 2 were

current annual growth of woody biomass < 1.4 m in

height and the number of pine trees > 8 cm dbh. In

addition to identifying species with affinities for

numerous large pines, this axis separated species

according to preference for woody understory biomass.

We were able to distinguish pine-grassland bird species

on CCA biplots by creating a pine-grassland variable by

combining large pine tree density and grass biomass

(Wilson 1994).

Forest interior species

Forest interior species tended to relate positively to

percent canopy cover, midstory foliage, overstory trees,

and tree species diversity. The black-and-white warbler

and ovenbird were strongly related to midstory foliage,

percent canopy cover, and oaks >8 cm dbh in the biplots

(Wilson 1994).. Scarlet tanager occurred infrequently,

but was clearly related to hardwoods. Summer tanager

related strongly to tree diversity in the floristic biplot

(Wilson 1994).
80 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Table 3.—Breeding bird responses (birds/40 ha)to wildlife stand improvement (WSI) and prescribed fire in pine-oak

stands on the Ouachita National Forest, Arkansas, May 1992, 1993, 1999 and 20001

 Treatment2 P > F

CONTROL WSI WSIB1 WSIB2 WSIB3 Contrasts3

Group, Species Mean(SE) Mean(SE) Mean(SE) Mean(SE) Mean(SE) Trt C vs WSI NB vs B

Forest Interior

Acadian Flycatcher 0.9(0.5) 0.6(0.4) 0.3(0.3) 1.3(0.9) 1.3(0.9) 0.485 0.232 0.666

American Redstart 0.0(0.0) 0.6(0.6) 0.5(0.5) 0.0(0.0) 0.0(0.0) 0.481 0.456 0.339

Black & White Warbler 13.3(3.6) a 9.2(2.4) ab 1.6(1.3) c 3.2(1.3) c 4.4(1.8) bc 0.009 0.010 0.012

Broad-winged Hawk 0.3(0.3) 1.6(1.3) 0.0(0.0) 0.3(0.3) 0.0(0.0) 0.444 0.734 0.129

Carolina Chickadee 8.8(3.4) 8.2(5.3) 10.7(4.1) 8.2(3.4) 3.8(1.7) 0.868 0.445 0.799

Cooper’s Hawk 0.0(0.0) 0.6(0.6) 0.0(0.0) 0.0(0.0) 0.3(0.3) 0.596 0.511 0.528

Downy Woodpecker 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.6(0.4) 0.9(0.9) 0.535 0.508 0.394

Great-crested Flycatcher 3.5(1.5) 8.5(3.4) 6.2(2.1) 10.7(3.0) 2.8(1.1) 0.135 0.215 0.764

Hairy Woodpecker 0.6(0.4) 1.9(1.0) 0.5(0.4) 0.0(0.0) 0.9(0.7) 0.421 0.692 0.166

Kentucky Warbler4 0.3(0.3) 0.6(0.6) 0.8(0.8) 0.6(0.6) 0.0(0.0) 0.903 0.588 0.705

Ovenbird 12.6(3.7) a 0.0(0.0) b 0.0(0.0) b 0.0(0.0) b 0.0(0.0) b <0.001 <0.001 1.000

Pileated Woodpecker 3.8(1.7) 2.2(1.1) 1.0(0.6) 0.9(0.7) 2.5(1.4) 0.656 0.264 0.347

Red-bellied Woodpecker 0.0(0.0) 0.0(0.0) 0.3(0.3) 0.3(0.3) 0.3(0.3) 0.397 0.225 0.124

Red-eyed Vireo 13.9(3.6) 16.7(3.7) 10.9(2.3) 16.7(4.0) 10.1(3.1) 0.512 0.931 0.258

Ruby-throated Hummingbird 0.3(0.3) 0.9(0.7) 1.3(0.7) 0.3(0.3) 1.3(0.9) 0.934 0.604 0.798

Scarlet Tanager 2.8(1.1) 1.6(1.6) 0.8(0.6) 1.9(1.9) 1.3(0.9) 0.193 0.023 0.895

Summer Tanager 8.5(2.3) 12.6(2.6) 9.6(2.9) 12.0(2.7) 10.1(2.7) 0.606 0.627 0.358

Tufted Titmouse 8.2(3.0) 4.1(1.6) 7.3(2.7) 0.9(0.9) 3.5(1.8) 0.063 0.076 0.652

Whip-poor-will4 0.9(0.7) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.414 0.061 1.000

Forest Edge and Other

American Crow 3.8(1.5) 0.3(0.3) 1.0(0.6) 1.3(1.3) 0.9(0.7) 0.221 0.033 0.582

American Goldfinch 2.2(1.4) c 4.7(1.9) bc 14.6(4.0) ab 12.0(3.5) ab 19.3(4.7) a 0.030 0.013 0.041

Blue Grosbeak 0.0(0.0) 0.0(0.0) 0.5(0.5) 0.0(0.0) 0.0(0.0) 0.460 0.651 0.559

Blue-gray Gnatcatcher 0.3(0.3) b 5.4(1.7) a 4.7(1.6) a 0.6(0.4) b 1.9(1.4) b 0.002 0.009 0.006

Blue Jay 5.4(2.3) 3.5(1.9) 1.3(1.1) 0.6(0.4) 1.9(1.3) 0.387 0.206 0.163

Brown-headed Cowbird 0.9(0.9) b 6.6(1.8) a 8.3(2.7) a 7.6(2.3) a 2.8(1.2) ab 0.041 0.009 0.439

Brown Thrasher 0.6(0.6) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.492 0.082 1.000

Catbird 0.0(0.0) 0.3(0.3) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.402 0.610 0.065

Carolina Wren 5.1(2.1) b 49.6(8.2) a 14.0(5.0) b 12.6(4.4) b 14.8(4.6) b 0.001 0.014 <0.001

Cedar Waxwing 0.0(0.0) 0.0(0.0) 0.3(0.3) 0.0(0.0) 0.0(0.0) 0.364 0.616 0.517

Common Flicker 0.0(0.0) 0.0(0.0) 1.3(0.7) 1.3(0.7) 0.6(0.6) 0.290 0.231 0.128

Common Grackle 0.0(0.0) 0.0(0.0) 0.3(0.3) 0.0(0.0) 0.0(0.0) 0.523 0.672 0.585

Common Yellowthroat 0.0(0.0) 0.9(0.9) 0.8(0.6) 3.2(1.5) 2.8(1.7) 0.119 0.060 0.204

Mourning Dove 0.0(0.0) 0.0(0.0) 1.6(1.1) 0.0(0.0) 0.6(0.6) 0.363 0.409 0.290

Northern Cardinal 1.3(0.9) 1.9(1.4) 0.8(0.6) 0.0(0.0) 1.3(1.3) 0.702 0.575 0.280

Red-shouldered Hawk 0.0(0.0) 0.0(0.0) 0.5(0.5) 0.0(0.0) 0.0(0.0) 0.460 0.651 0.559

Turkey Vulture 0.0(0.0) 0.6(0.4) 0.3(0.3) 0.0(0.0) 0.0(0.0) 0.497 0.552 0.107

White-breasted Nuthatch 0.3(0.3) 5.7(2.1) 5.2(2.0) 7.3(1.9) 3.5(1.7) 0.146 0.032 0.881

White-eyed Vireo 0.0(0.0) 0.3(0.3) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.402 0.610 0.065

Wild Turkey 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.6(0.6) 0.402 0.610 0.511

Yellow-billed Cuckoo 0.6(0.4) 2.8(1.9) 1.3(0.6) 2.8(1.4) 1.9(1.1) 0.811 0.324 0.653

Yellow-breasted Chat 0.0(0.0) c 2.2(1.2) bc 0.5(0.5) bc 4.7(2.2) ab 8.2(2.7) a 0.007 0.019 0.360

Yellow-throated Vireo 0.3(0.3) 0.0(0.0) 0.8(0.6) 0.9(0.7) 1.3(0.9) 0.852 0.833 0.293

Pine-grassland

Bachman’s Sparrow4 0.0(0.0) 0.6(0.6) 0.3(0.3) 1.9(1.0) 2.5(2.0) 0.217 0.125 0.525

Brown-headed Nuthatch4 0.0(0.0) b 0.0(0.0) b 10.1(4.0) a 6.3(2.7) a 4.4(2.6) ab 0.042 0.068 0.023

Continued
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 81

Chipping Sparrow 1.3(1.0) b 28.1(8.5) a 43.9(6.5) a 38.2(5.8) a 35.4(7.8) a 0.003 <0.001 0.071

Eastern Wood Pewee 0.0(0.0) c 11.1(3.3) ab 12.7(2.1) a 10.1(1.8) ab 6.0(2.2) b <0.001 <0.001 0.952

Indigo Bunting 4.7(2.3) c 57.2(9.2) a 36.4(4.5) b 54.0(6.7) a 64.1(7.3) a <0.001 <0.001 0.699

Northern Bobwhite 0.0(0.0) 2.8(1.3) 0.0(0.0) 0.6(0.6) 2.2(1.4) 0.098 0.156 0.064

Pine Warbler 61.3(8.1) 83.7(9.8) 93.4(7.3) 102.0(7.5) 96.6(7.3) 0.115 0.017 0.272

Prairie Warbler4 0.0(0.0) b 9.2(2.6) a 1.6(1.1) b 7.6(2.5) a 14.2(5.1) a 0.005 0.004 0.414

Red-cockaded Woodpecker4 0.0(0.0) b 0.0(0.0) b 1.8(1.0) a 2.2(1.4) a 0.9(0.7) a 0.048 0.053 0.017

Red-headed Woodpecker4 0.0(0.0) 3.2(2.2) 2.6(1.1) 2.5(1.4) 1.6(0.9) 0.183 0.022 0.776

1Column means within rows followed by the same letter or without letters were not significantly different at the 0.05 level (LSD). 2

Control=no treatment; WSI= midstory removal and no burning of downed logging slash or trees; WSIB1=midstory removal, late-winter

prescribed burn, following first growing season after fire; WSIB2=midstory removal, late-winter prescribed burn, following first growing

season after fire; WSIB3=midstory removal, late-winter prescribed burn, following first growing season after fire.

3

Specific orthogonal contrasts: C=control, WSI=midstory removal, NB=nonburned WSI stands, B=WSI stands burned within previous 3 years. 4

Priority species (Pashley et al. 2000).

Table 3.—continued

 Treatment2 P > F

CONTROL WSI WSIB1 WSIB2 WSIB3 Contrasts3

Group, Species Mean(SE) Mean(SE) Mean(SE) Mean(SE) Mean(SE) Trt C vs WSI NB vs B
82 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Carolina chickadee was positively related to tree

diversity, percent canopy cover, and midstory foliage in

the biplots (Wilson 1994). The pileated woodpecker was

positively related to percent canopy cover and large trees

in the CCA biplots (Wilson 1994). The red-eyed vireo,

tufted titmouse, and acadian flycatcher were positively

related to large hardwoods and tree diversity (Wilson

1994).

Forest edge species

Species associated with forest edge or canopy openings

were tolerant of wide ranges of canopy closure. These

species frequently occurred at intermediate levels of

midstory foliage and tree diversity. Blue jay was

positively related to percent canopy cover in the

physiognomy biplot. White-breasted nuthatch, yellowbilled cuckoo, and great-crested flycatcher occurred near

the center of both CCA biplots, indicating preferences

for intermediate values of physiognomy and floristic

measures (Wilson 1994). Carolina wren was positively

related to foliage 0-2 m and woody biomass <1.4 m, but

negatively related to pines >8 cm dbh. Wrens benefited

from the resulting slash from WSI (Wilson 1994).

The brown-headed cowbird was negatively related to

percent canopy cover, oaks >8 cm dbh, and midstory

foliage. Biplot positions for common yellowthroat and

yellow-breasted chat were greater than all other species

with respect to stems <1.4 m, foliage 0-2 m, and woody

biomass <1.4 m (Wilson 1994). American goldfinch

also favored open canopy structure (Wilson 1994).

Pine-grassland species

Pine-grassland species benefited most directly from

reduced canopy cover, reduced midstory, and associated

dense understory (Wilson 1994). We observed several

species associated with sites characterized by dense

understory and few trees. Prairie warbler and indigo

bunting demonstrated affinities for woody understory

vegetation with an aversion to overstory trees. These

species were negatively related to midstory foliage and

percent canopy cover. We found that pine warbler was

negatively related to tree diversity and midstory foliage,

and positively related to pines >8 cm dbh. Eastern wood

pewee appeared to key on scattered mature trees with an

open midstory. Pewees were negatively related to stems

>1.4 m, midstory foliage, tree diversity, and oaks >8 cm

dbh in CCA biplots (Wilson 1994).

The brown-headed nuthatch and chipping sparrow

demonstrated negative relationships to stems >1.4 m,

hardwoods >23.1 cm dbh, and tree diversity (Wilson

1994). Chipping sparrow was negatively associated with

stems <1.4 m and foliage 0-2 m. We found red-headed

woodpecker was negatively related to percent canopy

cover and midstory foliage and Bachman’s sparrow was

negatively related to percent canopy cover, midstory

foliage, oaks >8 cm dbh, and tree diversity (Wilson 1994).

Small Mammal Habitat Relationships

Most small mammal species were negatively correlated

with the presence of overstory and positively associated

with total biomass of understory production (Masters et

al. 1998). Total community abundance and whitefooted mouse abundance was positively associated with

dead woody debris and woody biomass in the

understory. However, total community abundance was

negatively associated with the presence of midstory and

more dense overstory (Masters et al. 1998). Both harvest

mice and golden mouse were positively associated with

the herbaceous component in the understory. As

herbaceous biomass increased so did both of these

species in addition to species richness and diversity

(Masters et al. 1998).

Discussion

Stand Characteristics

Forests of the Ouachita Mountains at settlement (ca.

1840 in Arkansas; ca. 1890 in Oklahoma) were

characterized by low-density open stands with scattered

large trees (Foti and Glenn 1991; Kreiter 1995). These

conditions were the result of the climatic (Foti and

Glenn 1991) and Native American fire regime ( Masters

et al. 1995). Stands currently undergoing restoration on

the Ouachita National Forest may have nearly twice the

stem density and one third more basal area as open pine

woodlands at settlement.

Wildlife stand improvement is an effective and timely

means of recreating open woodland structure because

periodic prescribed fire alone may take decades longer

to alter midstory structure in a similar fashion.

Following WSI, stands are characterized by pinedominated overstory, open midstory, and dense

understory of slash, hardwood sprouts, vines, grasses,

and forbs. The subsequent application of fire at 3-year

intervals appears to maintain this open midstory

structure, at least in the short-term, because fire has its

greatest impact on residual stems <8 cm dbh. Whereas

understory stem densities increase with time postburn,

fire top-kills small-diameter sprouts, maintaining an

open midstory. Understory structure and composition

will vary according to soils, slope, aspect, management

history, fire frequency and intensity, and time since

burned, thereby making available a range of habitats.

Bird Community

Bird community composition changed with alteration

of forest structure following WSI and fire (Figures 2 and

3). Analysis of bird species relationships with

physiognomy and floristic variables reveals species

habitat preferences across the broad continuum of

closed canopy pine-oak forest to open pine-grassland

woodlands. Several species were distinctly associated

with the woody structure that developed as plant
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 83

community succession progressed following fire. In

effect, a brief succession window of suitable habitat was

created for some species such as the common

yellowthroat and slightly longer windows for species

with broader habitat requirements (e.g., eastern wood

pewee, chipping sparrow, and indigo bunting) (Figure 3).

Seven species that occurred in our stands are listed as

priority species (Table 3; Pashley et al. 2000). However,

whip-poor-will was the only priority species that

showed even a tendency to be detrimentally affected by

restoration treatments in terms of frequency of

occurrence within the stands. The Kentucky warbler was

not affected. The remaining five priority species were

pine-grassland obligates and were distinctly benefited by

restoration treatments either in increased density or

frequency of occurrence (Tables 3 and 4).

Bird-habitat relationships

Forest interior species

Mature closed canopy forest habitats are necessary for

many interior forest species. Removal of midstory

structure followed by prescribed fire reduces densities

and diversity of midstory hardwoods, adversely affecting

species dependent on closed canopies or hardwood

midstory. Ground nesting species such as ovenbird and

black-and-white warbler may be adversely affected by

changes in understory particularly by the loss of the

litter layer following fire. Other forest interior species

may persist along riparian zones that dissect pinebluestem sites, but most will decline in thinned and

burned stands, particularly where hardwoods have

virtually been eliminated (Wilson et al. 1995). Thinning

and fire will not eliminate hardwood dependent species

such as red-eyed vireo if scattered mature hardwoods are

retained (Wilson et al. 1995).

Forest edge species

Tolerance of changes in forest structure varies among

forest edge species. Forest edge species of mixed forest

types (i.e., blue jay), will persist in stands retaining

hardwoods. Shrub-dwelling species such as Carolina

wren and yellow-breasted chat should occur in open

situations not recently burned. Whereas periodic fire

should benefit these species by promoting a shrubby

understory, annual burns may adversely affect habitat.

Ecotones created between restored pine-grassland

woodlands and closed canopy forest may provide

suitable habitats for some forest edge species.

Pine-grassland species

Occurrence and abundance of pine-grassland bird

species will vary with changes in understory. In addition

to red-cockaded woodpecker, we found brown-headed

nuthatch, Bachman’s sparrow, chipping sparrow, and

red-headed woodpecker are strongly associated with

habitat attributes of pine-grasslands. These species favor

open stands characterized by herbaceous understory and

sparse shrubs. Pine warbler and eastern wood pewee

also prefer open midstory, but may be more tolerant of

overstory hardwoods. Prescribed burning at periodic

intervals (3-5 years) will maintain forest structure

required by these species. As woody shrub layer increases

with time postburn, indigo bunting and prairie warbler

will increase. Fire regimes that eliminate woody shrubs

will exclude these two species.

Small Mammal Community

Pine-grassland restoration enhances the entire small

mammal community in terms of abundance, richness

and diversity. Without exception no species was

disadvantaged by restoration treatments. WSI and WSI

and fire created a more structurally complex understory

than in untreated stands. This was important for habitat

generalist species such as the white-footed mouse but

also for habitat specialist species such as the golden

mouse, harvest mouse, and the hispid cotton rat. The

later two species are notable because neither occurred in

control stands (Figure 4; Masters et al. 1998). Any

treatment that reduces overstory and allows for an

increase in understory production will likely benefit this

range of species. Fire likely played an important role

here as it reduced the stature (and thus shading effect)

of woody plants and fire reduced the litter layer,

promoting increases in herbaceous production (Masters

et al. 1998). Here again, the brief succession windows of

suitable habitat created by periodic fire were important

for some of the small mammal species (Figure 4).

Management Implications

Pine-grassland communities undoubtedly existed across

a broad continuum of canopy closures and basal areas

and should be viewed within a landscape-level context.

The landscape mosaic was likely composed of a range of

communities from open pine woodlands to more dense

hardwood-dominated sites such as mesic north slopes

and drainages (Foti and Glenn 1991). Breeding bird

habitat relationships reveal a need to provide variability

in stand structure and composition. This variability

should also enhance the range of small mammal species

in these stands. For both vertebrate communities, the

importance of periodic fire in creating brief windows of

suitable understory and midstory habitat structure for

some habitat specialist species cannot be

overemphasized. Upper and lower ends of targeted stem

densities and basal areas may be equally important in

maintaining representative habitats across the landscape.

Restoration of pine-bluestem habitats will be most

effective on sites of historical occurrence, including xeric

ridges and south slopes. In addition, the current practice

of maintaining hardwood corridors along stream

courses and drainages that dissect or encompass pinebluestem stands should be continued to benefit

numerous closed canopy and other species. We found
84 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Table 4.—Breeding bird responses (percent frequency of occurrence in bird plots) to wildlife stand improvement (WSI) and

prescribed fire in pine-oak stands on the Ouachita National Forest, Arkansas, May 1992, 1993, 1999 and 20001

Treatment2 P > F

CONTROL WSI WSIB1 WSIB2 WSIB3 Contrasts3

Group, Species Mean(SE) Mean(SE) Mean(SE) Mean(SE) Mean(SE) Trt C vs WSI NB vs B

Forest Interior

Acadian Flycatcher 4.8(2.3) 8.3(3.8) 2.6(1.3) 2.8(1.4) 3.2(1.3) 0.181 0.876 0.035

American Redstart 0.8(0.8) 6.0(2.9) 0.7(0.7) 1.6(1.1) 0.0(0.0) 0.160 0.528 0.027

Black & White Warbler 32.1(7.0) a 25.0(5.4) a 6.9(3.6) b 12.7(3.4) b 13.1(5.3) b 0.001 0.001 0.003

Broad-winged Hawk 3.2(1.3) 1.6(0.9) 1.0(0.7) 0.4(0.4) 0.8(0.8) 0.095 0.017 0.148

Carolina Chickadee 17.9(5.2) 16.7(4.0) 12.4(3.9) 13.1(4.4) 8.3(2.9) 0.626 0.332 0.282

Cooper’s Hawk 0.0(0.0) 0.8(0.8) 0.3(0.3) 0.0(0.0) 0.4(0.4) 0.697 0.414 0.693

Downy Woodpecker 0.8(0.8) 0.4(0.4) 0.3(0.3) 2.0(1.1) 2.0(1.4) 0.754 0.548 0.530

Great-crested Flycatcher 16.7(3.2) b 18.6(4.3) b 28.8(4.2) ab 39.3(5.7) a 18.3(3.4) b 0.022 0.087 0.072

Hairy Woodpecker 2.4(1.4) 4.0(1.6) 3.6(1.5) 1.6(1.1) 2.0(1.1) 0.735 0.924 0.420

Hooded Warbler 0.0(0.0) 0.0(0.0) 0.3(0.3) 0.0(0.0) 0.0(0.0) 0.523 0.672 0.585

Kentucky Warbler4 1.2(1.2) 2.4(1.3) 1.6(0.9) 2.4(1.7) 2.4(1.7) 0.954 0.547 0.641

Ovenbird 39.3(9.8) a 1.6(1.6) b 0.0(0.0) b 0.0(0.0) b 0.0(0.0) b <0.001 <0.001 0.328

Pileated Woodpecker 27.4(3.1) 24.6(4.6) 24.5(3.6) 20.2(3.0) 14.7(2.8) 0.063 0.038 0.435

Red-bellied Woodpecker 0.4(0.4) 2.0(1.1) 3.6(1.3) 1.2(1.2) 4.8(1.9) 0.117 0.124 0.422

Red-eyed Vireo 58.3(7.5) 56.4(4.1) 45.1(5.8) 54.8(5.5) 40.1(6.3) 0.073 0.057 0.128

Ruby-throated Hummingbird 0.4(0.4) 1.2(0.9) 1.6(0.9) 0.4(0.4) 1.6(1.1) 0.934 0.604 0.798

Scarlet Tanager 16.7(5.1) 10.7(2.8) 7.2(1.8) 4.8(3.2) 8.7(4.0) 0.130 0.082 0.174

Summer Tanager 30.6(5.6) 26.6(4.3) 37.6(3.9) 40.1(5.7) 28.6(4.9) 0.228 0.832 0.070

Tufted Titmouse 39.7(6.6) 27.8(4.4) 28.4(4.3) 19.8(7.4) 2 3.0(6.7) 0.115 0.037 0.395

Whip-poor-will4 0.8(0.5) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.414 0.061 1.000

Wood Thrush 2.4(2.4) 0.0(0.0) 0.7(0.7) 0.0(0.0) 0.4(0.4) 0.722 0.645 0.454

Worm-eating Warbler 0.0(0.0) 0.0(0.0) 1.0(1.0) 0.0(0.0) 0.0(0.0) 0.523 0.672 0.585

Forest Edge and Other

American Crow 25.8(4.0) 32.1(5.8) 30.7(5.4) 34.1(5.9) 32.9(6.8) 0.892 0.356 0.915

American Goldfinch 9.9(3.4) b 11.5(3.9) b 22.5(3.6) a 23.0(5.3) a 26.6(6.4) a 0.024 0.020 0.010

American Kestrel 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.4(0.4) 0.402 0.610 0.511

Barred Owl 1.2(0.9) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.063 0.005 1.000

Blue Grosbeak 0.0(0.0) 2.0(1.1) 1.0(0.7) 0.0(0.0) 0.8(0.8) 0.061 0.115 0.021

Blue-gray Gnatcatcher 0.4(0.4) b 8.3(2.7) a 6.9(1.8) a 1.6(0.9) b 3.6(2.8) b 0.003 0.006 0.017

Blue Jay 24.2(5.2) 11.9(3.5) 6.2(2.1) 6.3(1.9) 8.7(3.2) 0.083 0.010 0.302

Brown-headed Cowbird 1.2(0.9) b 17.1(2.7) a 21.6(4.8) a 20.6(3.9) a 11.5(3.3) a <0.001 <0.001 0.546

Brown Thrasher 1.6(1.6) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.492 0.082 1.000

Catbird 0.0(0.0) 0.4(0.4) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.402 0.610 0.065

Carolina Wren 21.8(4.7) c 79.0(4.7) a 51.6(7.2) b 45.6(6.9) b 46.4(5.6) b <0.001 0.001 <0.001

Cedar Waxwing 0.0(0.0) 0.0(0.0) 1.2(1.2) 0.9(0.9) 0.0(0.0) 0.700 0.587 0.487

Chimney Swift 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.6(0.6) 0.0(0.0) 0.450 0.627 0.531

Common Flicker 0.8(0.5) 2.0(1.1) 7.5(2.2) 2.8(1.1) 5.2(2.1) 0.202 0.122 0.146

Common Grackle 0.0(0.0) 0.0(0.0) 0.4(0.4) 0.0(0.0) 0.0(0.0) 0.519 0.676 0.589

Common Yellowthroat 0.0(0.0) c 5.2(3.5) bc 4.2(2.3) ab 13.1(5.8) a 17.1(8.2) ab 0.011 0.004 0.068

Eastern Bluebird 0.0(0.0) 0.8(0.5) 3.9(2.6) 0.8(0.8) 0.4(0.4) 0.210 0.116 0.687

Field Sparrow 0.0(0.0) 1.6(1.6) 0.0(0.0) 0.0(0.0) 0.8(0.8) 0.488 0.454 0.324

Fish Crow 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.4(0.4) 0.414 0.615 0.516

Great-horned Owl 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.4(0.4) 0.4(0.4) 0.397 0.410 0.291

Mourning Dove 0.0(0.0) 4.4(1.9) 5.9(2.9) 5.6(3.3) 6.3(3.8) 0.233 0.031 0.764

Northern Cardinal 6.0(2.2) 9.5(3.4) 4.2(2.0) 0.8(0.8) 4.4(3.2) 0.287 0.569 0.062

Northern Parula 0.0(0.0) 0.0(0.0) 0.3(0.3) 0.0(0.0) 0.0(0.0) 0.523 0.672 0.585

Continued
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 85

Purple Martin 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.9(0.9) 0.500 0.644 0.553

Red-shouldered Hawk 2.8(1.5) 0.8(0.8) 3.3(2.3) 0.8(0.8) 0.0(0.0) 0.494 0.142 0.915

Red-tailed Hawk 0.4(0.4) 0.0(0.0) 2.0(1.2) 1.2(0.9) 0.4(0.4) 0.340 0.779 0.085

Screech Owl 0.4(0.4) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.402 0.058 1.000

Turkey Vulture 0.0(0.0) 1.6(1.1) 0.3(0.3) 0.0(0.0) 0.0(0.0) 0.493 0.553 0.104

White-breasted Nuthatch 2.0(1.1) b 19.0(4.5) a 20.6(2.8) a 25.4(4.9) a 15.9(4.0) a 0.024 0.003 0.664

White-eyed Vireo 0.0(0.0) 2.8(1.5) 0.0(0.0) 1.6(1.1) 0.0(0.0) 0.127 0.274 0.043

Wild Turkey 2.4(2.4) 0.8(0.8) 4.2(2.7) 7.9(2.5) 4.0(1.8) 0.100 0.152 0.055

Wood Duck 0.0(0.0) 0.0(0.0) 0.6(0.6) 0.9(0.9) 0.0(0.0) 0.673 0.587 0.488

Yellow-billed Cuckoo 17.1(4.8) 18.3(4.9) 7.8(2.3) 13.9(4.9) 14.7(4.5) 0.352 0.428 0.205

Yellow-breasted Chat 3.2(1.8) b 29.4(8.0) a 10.1(2.9) b 34.1(8.9) a 43.3(8.1) a <0.001 <0.001 0.882

Yellow-throated Vireo 0.4(0.4) 6.0(3.2) 8.5(2.7) 4.8(2.2) 6.7(2.7) 0.342 0.092 0.543

Pine-grassland

Bachman’s Sparrow4 0.0(0.0) c 2.0(2.0) c 4.6(2.9) bc 11.1(4.0) ab 13.5(4.8) a 0.019 0.023 0.037

Brown-headed Nuthatch4 0.0(0.0) c 0.8(0.8) bc 7.8(3.3) ab 8.7(3.7) a 6.3(2.3) ab 0.030 0.025 0.017

Chipping Sparrow 3.2(1.8) c 36.5(8.0) b 69.0(4.9) a 55.6(6.5) ab 55.6(6.9) ab <0.001 <0.001 0.010

Eastern Wood Pewee 0.8(0.8) c 38.5(5.1) b 53.3(4.0) a 40.1(4.3) ab 32.9(6.5) b <0.001 <0.001 0.371

Indigo Bunting 19.4(5.3) b 73.4(4.3) a 69.6(4.9) a 78.2(4.3) a 77.0(5.0) a <0.001 <0.001 0.582

Northern Bobwhite 2.8(1.7) c 34.1(6.7) a 14.4(3.7) b 15.5(6.4) bc 32.1(9.9) ab 0.011 0.005 0.017

Pine Warbler 94.8(1.7) 92.5(2.6) 95.8(1.9) 96.0(1.4) 9 5.2(1.9) 0.504 0.745 0.144

Prairie Warbler4 1.2(0.9) c 27.8(7.1) ab 13.4(4.1) bc 39.3(8.0) a 48.0(8.6) a 0.002 <0.001 0.291

Red-cockaded Woodpecker4 0.0(0.0) b 0.0(0.0) b 3.6(1.7) a 2.4(1.1) a 3.2(1.8) a 0.014 0.020 0.005

Red-headed Woodpecker4 0.0(0.0) b 7.5(4.1) ab 14.7(3.3) a 9.1(4.2) ab 5.6(2.7) ab 0.045 0.015 0.335

1

Column means within rows followed by the same letter or without letters were not significantly different at the 0.05 level (LSD). 2

Control=no treatment; WSI= midstory removal and no burning of downed logging slash or trees; WSIB1=midstory removal, late-winter

prescribed burn, following first growing season after fire; WSIB2=midstory removal, late-winter prescribed burn, following first growing

season after fire; WSIB3=midstory removal, late-winter prescribed burn, following first growing season after fire.

3

Specific orthogonal contrasts: C=control, WSI=midstory removal, NB=nonburned WSI stands, B=WSI stands burned within previous 3 years. 4

Priority species (Pashley et al. 2000).

Table 4.—continued

Treatment2 P > F

CONTROL WSI WSIB1 WSIB2 WSIB3 Contrasts3

Group, Species Mean(SE) Mean(SE) Mean(SE) Mean(SE) Mean(SE) Trt C vs WSI NB vs B
86 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Carolina wren, Kentucky warbler, red-eyed vireo and

tufted titmouse were closely associated with greenbelts

or drainages dominated by hardwoods extending into

the interior of treated stands. Residual hardwoods

retained within the stand were important to red-eyed

vireo, summer tanager, scarlet tanager, tufted titmouse,

yellow-throated vireo and white-breasted nuthatch

occurrence within a restored stand.

Although relative abundance and frequency of

occurrence increased for many species, we recognize that

this does not necessarily translate to high reproductive

success. Nest predation and parasitism can be a major

concern in pine forests of the southeast U.S. (White et

al. 1999). A concern is raised here because of increased

brown-headed cowbird abundance in stands that had

been recently thinned (WSI) and in the first and second

growing season following fire (WSIB1 and WSIB2

treatments respectively). We did observe fledglings

during point-counts of brown-headed nuthatch,

Carolina wren, chipping sparrow, common

yellowthroat, indigo bunting, ovenbird and pine warbler

in stands associated with their highest densities, but not

all observers noted occurrence of fledglings. The

influence of cowbird nest parasitism in restored stands

should be further examined.

While knowledge of individual species relationships to

habitat components is vital to manage for habitat

availability (Wilson et al. 1995, Masters et al. 1998),

regional biodiversity will likely be maximized by

restoring the natural disturbance regime (Vose 2000). By

mimicking fire events of the past, the most complete

range of habitats and attendant species may best be

maintained over time in the Ouachita Mountains

ecosystem and possibly shortleaf pine dominated

systems elsewhere. With repeated alternating shifts from

frequent burning (<3 year intervals) to no burning and

periodic shifts to growing season burns, current stands

may eventually approach structural conditions reported

historically as they are repeatedly thinned. Irregular

burn intervals may be particularly important to ensure

adequate pine recruitment (Cain 1993; Masters et al

1995). A return to the presettlement fire regime will not

eliminate residual hardwoods because large hardwoods

are tolerant even of frequent fire (Waldrop et al. 1992).

Plant and animal response to large-scale burns and

growing season burns should be investigated as such

burns were probably characteristic of the original fire

regime (Masters et al 1995).

Landscape-level studies are necessary to determine avian

relationships to stand sizes and juxtaposition of pinebluestem woodlands and pine-oak forest. Geographic

information systems and ecological models should be

used to ensure the presence of uncommon natural

communities across the landscape and into the future.

We recommend additional research on red-cockaded

woodpecker population viability in low-density stands

with varying proportions of hardwoods because overall

stand density may be as critical as some upper level

threshold for a maximum hardwood component.

Research is also needed to determine the extent and

importance of other fire-dependent community types,

including oak-dominated woodlands and savannas and

breeding bird and small mammal habitat relationships

within those settings.

Acknowledgments

Funding was provided by the USDA Forest Service and

Oklahoma State University Forestry Department and inkind support was provided by Arkansas Game and Fish

Commission. We thank J. Neal for assistance with

breeding bird surveys and W. Montague for assistance

with identifying stands and coordination of burning

efforts. R. Putnam, S. Hooks, H. Johnson, J. Ridenhour,

E. Stewart, R. Stewart, K. Piles, and C. Brown are

gratefully acknowledged for assistance with field work.

We are grateful to M. Payton for assistance with

experimental design and statistical analysis and S.

Ditchkoff for assistance with analysis and graphics. This

publication is approved by the Director of the Oklahoma

State University Agricultural Experiment Station.

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List of Common and Scientific Names of all Occurring Breeding Bird and Small

Mammal Species

Common Name Scientific Name (authority)

Birds

Acadian Flycatcher Empidonax virescens (Vieillot)

American Crow Corvus brachyrhynchos (Brehm)

American Goldfinch Carduelis tristis (Linnaeus)

American Kestrel Falco sparverius (Linnaeus)

American Redstart Setophaga ruticilla (Linnaeus)

Bachman’s Sparrow Aimophila aestivalis (Lichtenstein)

Barred Owl Strix varia (Barton)

Black and White Warbler Mniotilta varia (Linnaeus)

Blue Grosbeak Guiraca caerulea (Linnaeus)

Blue Jay Cyanocitta cristata (Linnaeus)

Blue-gray Gnatcatcher Polioptila caerulea (Linnaeus)

Broad-winged Hawk Buteo platypterus (Vieillot)

Brown Thrasher Toxostoma rufum (Linnaeus)

Brown-headed Cowbird Molothrus ater (Boddaert)

Brown-headed Nuthatch Sitta pusilla (Latham)

Carolina Chickadee Poecile carolinensis (Audubon)

Carolina Wren Thryothorus ludovicianus (Latham)

Catbird Dumetella carolinensis (Linnaeus)

Cedar Waxwing Bombycilla cedrorum (Vieillot)

Chimney Swift Chaetura pelagica (Linnaeus)

Chipping Sparrow Spizella passerina (Bechstein)

Common Flicker Colaptes auratus (Linnaeus)

Common Grackle Quiscalus quiscula (Linnaeus)

Common Yellowthroat Geothlypis trichas (Linnaeus)

Cooper’s Hawk Geothlypis trichas (Bonaparte)

Downy Woodpecker Picoides pubescens (Linnaeus)

Eastern Bluebird Sialia sialis (Linnaeus)

Eastern Wood Pewee Contopus virens (Linnaeus)

Field Sparrow Spizella pusilla (Wilson)

Fish Crow Corvus ossifragus (Wilson)

Great-crested Flycatcher Myiarchus crinitus (Linnaeus)

Great-horned Owl Bubo virginianus (Gmelin)

Hairy Woodpecker Picoides villosus (Linnaeus)

Hooded Warbler Wilsonia citrina (Boddaert)

Indigo Bunting Passerina cyanea (Linnaeus)

Kentucky Warbler Oporornis formosus (Wilson)

Mourning Dove Zenaida macroura (Linnaeus)

Northern Bobwhite Colinus virginianus (Linnaeus)

Northern Cardinal Cardinalis cardinalis (Linnaeus)

Northern Parula Parula americana (Linnaeus)

Ovenbird Seiurus aurocapillus (Linnaeus)

Pileated Woodpecker Dryocopus pileatus (Linnaeus)

Pine Warbler Dendroica pinus (Wilson)

Prairie Warbler Dendroica discolor (Vieillot)

Purple Martin Progne subis (Linnaeus)

Red-bellied Woodpecker Melanerpes carolinus (Linnaeus)

Red-cockaded Woodpecker Picoides borealis (Vieillot)

continued
90 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Red-eyed Vireo Vireo olivaceus (Linnaeus)

Red-headed Woodpecker Melanerpes erythrocephalus (Linnaeus)

Red-shouldered Hawk Buteo lineatus (Gmelin)

Red-tailed Hawk Buteo jamaicensis (Gmelin)

Ruby-throated Hummingbird Archilochus colubris (Linnaeus)

Scarlet Tanager Piranga olivacea (Gmelin)

Screech Owl Otus asio (Linnaeus)

Summer Tanager Piranga rubra (Linnaeus)

Tufted Titmouse Baeolophus bicolor (Linnaeus)

Turkey Vulture Cathartes aura (Linnaeus)

Whip-poor-will Caprimulgus vociferous (Wilson)

White-breasted Nuthatch Sitta carolinensis (Latham)

White-eyed Vireo Vireo griseus (Boddaert)

Wild Turkey Meleagris gallopavo (Linnaeus)

Wood Duck Aix sponsa (Linnaeus)

Wood Thrush Hylocichla mustelina (Gmelin)

Worm-eating Warbler Helmitheros vermivorus (Gmelin)

Yellow-billed Cuckoo Coccyzus americanus (Linnaeus)

Yellow-breasted Chat Icteria virens (Linnaeus)

Yellow-throated Vireo Vireo flavifrons (Vieillot)

Small Mammals

Cotton Mouse Peromyscus gossypinus (LeConte)

Deer Mouse Peromyscus maniculatus (Wagner)

Eastern Woodrat Neotoma floridana (Ord)

Fulvous Harvest Mouse Reithrodontomys fulvescens (J.A. Allen)

Hispid Cotton Rat Sigmodon hispidus (Say and Ord)

Golden Mouse Ochrotomys nutalli (Harlan)

Pine Vole Microtus pinetorum (LeConte)

Southern Short-tailed Shrew Blarina carolinensis (Bachman)

White-footed Mouse Peromyscus leucopus (Rafinesque)

List of Common and Scientific Names—continued

Common Name Scientific Name (authority)
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 91

Abstract.—In this paper I review studies of herpetofauna

in two fire-maintained, xeric pineland southeastern U.S.

ecosystems, longleaf pine-turkey oak sandhills and

Florida sand pine scrub. I address evolutionary

adaptations of herpetofauna to these xeric

environments, and how fire disturbance influences

herpetofaunal community composition by structuring

habitat. Where data are available I examine how

anthropogenic disturbance such as fire suppression,

clearcutting, and restoration treatments can affect

herpetofaunal community composition, and whether

some anthropogenic disturbance types can mimic

natural disturbance in their effects. I also address

possible reasons why detection of population response

is more difficult for amphibians than for common

reptile species.

Natural disturbance is an important force in shaping the

habitat structure of ecosystems within southeastern

United States. However, type, frequency, intensity, and

scale of natural disturbance varies among ecosystems.

Wind, ice, insect and disease outbreaks, or fire can be

dominant disturbance types in some southeastern

ecosystems, acting at scales ranging from single-tree

death to thousands of hectares. Disturbance regimes and

the habitat structure they maintain exert selective

pressure on life history strategies, physical, and

behavioral adaptations of resident flora and fauna

(Denslow 1980). Hence, biotic components that

characterize and distinguish ecosystems often provide

clues to the most common habitat structure, and

dominant disturbance regime that historically shaped it.

Interference with disturbance regimes can alter the

habitat structure of ecosystems by disrupting processes

that shape, maintain, and perpetuate them. Major

structural alterations to habitat can, in turn, potentially

adversely affect characteristic flora and fauna. Similarly,

introduction of anthropogenic disturbance types to

ecosystems that do not resemble or mimic the results of

natural disturbance regimes can adversely affect their

biota.

Fire is an integral disturbance type to many southeastern

ecosystems that drives their process, function, and many

of their defining floral and faunal components (Myers

and Van Lear 1998). However, fire characteristics vary

among ecosystems. In the southeastern Coastal Plain,

the longleaf pine-wiregrass ecosystem was historically

maintained by low-intensity, high frequency (3-5 year

intervals), large-scale groundfires that killed many

young, invading hardwoods. This burning regime

maintained a low density of mature longleaf pine,

scattered understory oaks, and an abundance of sunlight

at ground level that promoted a continuous

groundcover dominated by wiregrass and a high

diversity of herbaceous plants (Myers 1990). Fire

suppression is more aptly viewed as a disturbance in

sandhills than is fire itself, as it disrupts a process that

maintains and perpetuates this habitat structure. In the

absence of fire a dense hardwood understory develops,

leaf litter and shade increase, and wiregrass becomes

sparse (see Table 1).

Conversely, the sand pine scrub ecosystem in peninsular

Florida was maintained by high-intensity, low frequency

(10-100 year intervals) wildfires that killed virtually all

above-ground vegetation, followed by rapid resprouting

of shrubs (Abrahamson 1984; Greenberg et al. 1995).

During long fire-free intervals a thick shrub layer

developed, and even-aged stands of sand pine matured

at densities that varied dramatically among scrubs.

Herpetofaunal assemblages differ among southeastern

ecosystems and physiographical regions according to

historical biogeography, and physiological and

environmental constraints that adapt each species to

specific or generalized habitats. Species exhibiting

specialized traits often are also restricted in their local

distribution by habitat type and condition. Reptile and

amphibian species in xeric sandhills and scrub include

widespread generalists and species having narrow

habitat requirements, usually typical of recent post-fire

conditions. Because the habitat condition in xeric

sandhills and sand pine scrub is sculpted by fire,

herpetofaunal distribution and abundance is closely tied

to disturbance regime.

Surprisingly few studies examine differences in habitat

characteristics between fire-suppressed and firemaintained sandhills or sand pine scrub, and how

habitat structure affects herpetofauna. However, even an

intuitive understanding of fire effects and ecological

requirements of herpetofauna would suggest that there

are impacts. Species having specialized adaptations to

structural features of fire-maintained habitat, such as

bare ground, could be at a disadvantage in fire

suppressed habitat. Gopher tortoises are abundant in

regularly burned sandhills, where a dense layer of

wiregrass and herbaceous groundcover provide plentiful

food, and the light level is adequate for nest sites

(Diemer 1992). This might be expected to adversely

affect gopher tortoise burrow commensals, but only if

burrows are a limiting resource. By reducing root density

in uplands (by reducing tree density), fire likely

Fire, Habitat Structure and Herpetofauna in the Southeast

Cathryn H. Greenberg1

1

Southern Research Station, USDA Forest Service, Asheville,

NC 28806.
92 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

facilitates burrowing conditions for primary

underground excavators (such as gopher tortoises and

pocket gophers) and thereby secondary burrow- and

tunnel users such as Florida gopher frogs and striped

newts. Historically, fire probably maintained and

deepened ephemeral wetlands that are used as breeding

sites by xeric pineland amphibians by killing

encroaching trees and shrubs, and burning peat and

vegetation buildup during dry periods. Fire also

maintained a low density of mature trees in uplands

surrounding ponds by killing young trees. Lower tree

density reduces transpiration, but may increase

evaporation from ponds. Hardwood invasion may affect

hydroperiod and water depth of shallow ponds by

altering the balance between transpiration and

evaporation (Sharitz and Gresham 1998).

In this paper I will review studies of herpetofauna in two

fire-maintained xeric pineland ecosystems, longleaf

pine-turkey oak-wiregrass sandhills and Florida sand

pine scrub. I will specifically address some evolutionary

adaptations of herpetofauna to xeric environments of

these fire-maintained ecosystems, and how disturbance

influences herpetofaunal community composition by

structuring habitat. I also will discuss possible reasons

why detection of population response is more difficult

for amphibians than for common reptile species. Where

data are available I examine how anthropogenic

disturbance such as fire suppression, clearcutting, and

restoration treatments can affect herpetofaunal

community composition, and whether some

anthropogenic disturbance types can mimic natural

disturbance in their effects.

Although reptiles and amphibians often are generically

lumped together as “herpetofauna,” they are

phylogenetically as distinct from one another as are

mammals from birds. Amphibians (class Amphibia) have

permeable, moist skin that for many serves a respiratory

function, and increases their susceptibility to

desiccation. Many species also require water for egg

deposition and larval development. Taxa vary

considerably in their vulnerability to desiccation. For

example, anurans are more tolerant of high

temperatures (Stebbins and Cohen 1995), and can store

and reabsorb larger amounts of water in their bladders

than salamanders (Zug 1993). Some salamanders are

lungless, and some are completely terrestrial

(DeMaynadier and Hunter 1995). Many amphibian

species have small home ranges (Duellman and Trueb

1986) and poor dispersal capabilities (Sinsch 1990).

Conversely, most reptiles (class Reptilia) require warm

temperatures (associated with higher light levels) for egg

incubation and successful development of hatchlings

(Deeming and Ferguson 1991). Reptiles have dry, scaly

skin that protects them from desiccation. Clearly,

response to disturbance might be expected to differ

between the two classes, and among species within

them.

Behavioral or physical adaptations to xeric upland

environments likely reduce the vulnerability of the

native herpetofauna to fire and consequent habitat

alterations (Means and Campbell 1981). Among

reptilian sandhill and sand pine scrub inhabitants,

several species exhibit specialized behavioral or

anatomical traits for surviving in a xeric, open, loosesand environment (Campbell and Christman 1982a;

Table 1.—Mean (+SE) percent cover of select habitat characteristics and basal area

(BA) of trees in hardwood-invaded (HI) versus savanna-like (SL) longleaf pinewiregrass sandhills surrounding eight isolated, ephemeral ponds in the Ocala

National Forest, Florida. Percentage data were square root-arcsine transformed for

t-tests but are presented as untransformed values (adapted from Greenberg in press)

 Upland matrix

HI SL df P

Wiregrass (% cover) 11.6+ 6.0 66.5+8.6 25.0 <0.0001

Herbaceous (% cover) 4.5 + 1.4 21.5+4.7 25.0 0.0002

Leaf litter (% cover) 99.3+ 0.5 68.5+9.6 12.7 0.0023

Bare ground (% cover) 0.4 + 0.2 9.4+3.7 13.3 0.0072

Coarse woody debris 0.7 + 0.4 1.2+0.6 25.0 0.6208

>12.5 cm (% cover)

Shrub <2.5 cm (% cover) 47.5+8.2 27.8+6.4 25.0 0.0548

Light (%) 22.8+3.7 54.0+4.9 25.0 <0.0001

Longleaf pine BA (m2

) 4.5 +1.2 4.8+1.1 25.0 0.8475

Hardwoods & sand pine BA (m2

) 2.8+0.7 0.9+0.5 25.0 0.0357

Snag BA (m2

) 0.7+0.3 0.5+0.3 25.0 0.5677
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 93

Greenberg et al. 1994; Stout et al. 1998). For example,

sand skinks, mole skinks, and crown snakes are “sand

swimmers,” requiring loose, bare sand for semifossorial

movement. Short-tailed snakes (Campbell 1992) and

pine snakes (Conant and Collins 1991) use

underground burrows extensively. Six-lined racerunners

and scrub lizards are cursorial, and the mechanics of

movement are apparently facilitated by open, bare sand.

Behaviors that exploit cool, moist microhabitats also are

apparent in many reptile species inhabiting sandhills

and scrub. Gopher tortoises dig burrows that are used as

retreats by at least 23 reptile species, including pine

snakes and sand skinks, and 9 amphibian species,

including gopher frogs and narrowmouth toads

(Jackson and Milstrey 1989). Several species, including

southern, eastern narrowmouth, and spadefoot toads

avoid heat and desiccation by burrowing underground,

or by using pre-existing burrows, tunnels and root

channels (Russell et al. 1999). Nocturnal foraging, that

may allow some species to tolerate extreme conditions

of temperature and drought. Selective pressure “preadapts” many species to their environments and the

disturbance regimes that shape them.

Many reptile species appear to respond to firemaintained habitat features such as the amount of bare

ground, rather than to plant associations per se

(Campbell and Christman 1982; Greenberg et al. 1994).

In sand pine scrub, scrub lizards, sand skinks, six-lined

racerunners, mole skinks, and crown snakes are more

abundant in open scrub habitat with a high proportion

of bare sand than in mature, forested scrub (Campbell

and Christman 1982; Greenberg et al. 1994; Means and

Campbell 1982). Scrub lizards (r2

 = 0.44; p = 0.0182)

and six-lined racerunners (r2

 = 68; p = 0.0010) are

positively correlated, whereas southeastern five-lined

skinks are negatively correlated (r2 = 0.75; p = 0.0003)

to the amount of bare sand (Greenberg et al.1994).

Reptile response to anthropogenic and natural

disturbances may be similar in some ecosystems if the

resulting post-disturbance habitat structure is similar.

For example, the relative abundance of mole skinks,

scrub lizards, and six-lined racerunners was similar in

sites that had been recently burned and salvage-logged,

clearcut with low-intensity site preparation, and clearcut

followed by roller-chopping, but lower in mature

forested sand pine scrub (Greenberg et al. 1994).

Conversely, generalist species that are common in many

southeastern ecosystems, such as ground skinks and

southeastern five-lined skinks were widespread among

scrub age classes or more abundant in mature sand pine

forest than in young, open scrub (Table 2) (Greenberg et

al. 1994). This suggests that high-intensity natural

disturbance was an important selective pressure on

physical or behavioral adaptations by herpetofauna.

There is a dearth of information on how fire

suppression affects herpetofauna of xeric sandhills.

Potential effects appear difficult to detect, perhaps in

part because habitat heterogeneity remains within

hardwood-invaded sandhills for decades after

commencement of fire suppression. Litt (1999) reported

no difference in species richness, and few differences in

relative abundance of species between fire-suppressed

sandhills and three restoration treatments (mechanical

girdling and felling; herbicide; and growing season fire)

or frequently burned “reference” sandhills. Meshaka and

Layne (pers. comm.) also found that during 1979-1994,

species diversity and richness of herpetofauna in a firesuppressed sandhill in Florida was similar to that

reported for frequently burned Florida sandhills. They

attributed the persistence of xeric-adapted reptiles such

as six-lined racerunners and scrub lizards species to a

frequent occurrence of shrub-free openings. The

abundance of southeastern five-lined skinks increased

during their study. Mushinsky (1985) reported higher

abundance of six-lined racerunners on an annually

burned sandhill site with low wiregrass and herbaceous

cover relative to sites with longer burn intervals. Sandhill

reptiles respond to subsets of microhabitat

characteristics that are unique to each species or species

group (Litt 1999; Mushinsky 1985). Habitat

Table 2.—Mean (+ SE) number of select reptile species captured using drift fences and pitfall traps during

August 1991 - September 1992 in three treatments and mature forest controls (n = 3 per treatment and

control) in sand pine scrub, Ocala National Forest, Florida

Species Burn & Clearcut & Clearcut & Mature P

Salvage Rollerchop Bracke-Seed Forest

Six-lined Racerunner 5.7 + 1.2a,b 12.3 + 1.2a 13.0 + 5.2a 2.3 + 1.5b 0.0752

Mole Skink 11.0 + 2.1a 16.7 + 3.5a 9.7 + 3.5a 1.3 + 0.3b 0.0237

Southeastern Five-lined Skink 1.3 + 1.3a 1.0 + 0.6a 1.0 + 0.6a 13.3 + 1.7b 0.0001

Ground Skink 2.0 + 1.0a,b 0.7 + 0.7a 0.0 + 0.0 3.7 + 0.3b 0.0140

Scrub Lizard 12.0 + 6.5a,b 25.0 + 11.0a,c 38.3 + 2.6c 1.7 + 0.3b 0.0194

Peninsula crown snake 2.0 + 1.2a 7.7 + 2.4a,b 8.7 + 1.3b 3.7 + 2.2a 0.0977
94 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

heterogeneity within regularly burned sandhills permits

the co-occurrence of post-disturbance specialists and

generalists, and contributes to the high diversity of

herpetofauna in sandhills and scrub relative to other

Florida ecosystems (Campbell and Christman 1982).

Among the few studies comparing amphibian use of fire

suppressed versus regularly burned xeric pinelands,

none provide conclusive results. Clearly, amphibians

that are characteristic of xeric, fire-climax pinelands also

use fire-suppressed habitat (Greenberg 2001; Greenberg

unpubl. data; Litt 1999; Mushinsky 1985). Palis (1998)

reported high adult usage and egg mass deposition by

gopher frogs at a pond within fire-suppressed longleaf

pine-turkey oak sandhill uplands. Some studies provide

anecdotal evidence of habitat preference. Palis (1998)

reported a tendency for adult gopher frogs to immigrate

to a breeding pond from the direction of an early

successional bombing range versus a hardwood-invaded

sandhill, suggesting heavier use of the more open

habitat. Dodd (1996) reported more captures in open

xeric hammock, fewer in closed xeric hammock, and as

expected in sandhills if captures were proportional to

trap effort.

Season of burn influences diversity patterns, percent

cover, and flowering of several herbaceous sandhill

species (Robins and Myers 1989). Few studies address

whether season of burn affects herpetofaunal

communities in sandhills. C.L. Hardy and associates

(pers. comm.) applied dormant season burns to uplands

surrounding one half of ephemeral ponds in sandhills,

and growing season burns to the other half. Their

preliminary results indicate no treatment differences in

amphibian or reptile use of ponds.

Amphibian population response to disturbance or

habitat condition appears to be difficult to detect in fireadapted ecosystems such as sandhills and sand pine

scrub, and may require long-term data. Although many

species use uplands during much of their lives, the

distribution and abundance of amphibians in uplands is

closely tied to distance from water (Dodd 1995, 1996).

Greenberg (1993) reported that the relative abundance

of anurans was not significantly related to stand age or

disturbance treatment in sand pine scrub, but was

negatively correlated with distance from permanent

water sources (lakes and wet prairies) (r2

 = 0.5406; p =

0.0064). This relationship was not consistent among

individual species, possibly because some small,

ephemeral wetlands were not detected, and because

some species travel further from breeding sites than

others (Dodd 1996). Dodd (1996) reported no

significant correlation between the number of

amphibians captured per trap and trap distance from

nearest water body, but 83% of amphibians were

captured within 600-m from water. Greenberg (1993)

collected spadefoot toad tadpoles from a roadside

puddle in sand pine scrub. This suggests larval

developmental rates, precipitation, and the distribution

of depressions as small as puddles (in this case,

anthropogenically-created by clay-subsidized sand

roads) can differentially affect the spatial and temporal

distribution patterns of amphibians. Clearly, ephemeral

ponds are critical centers of herpetofaunal diversity in

sandhills (Dodd 1992; Moler and Franz 1987); most

amphibian and several reptile species, such as turtles

and swamp snakes, would not occur there in their

absence.

Wetlands within sandhills function as a magnet to

amphibians. Capture rates may reflect life history

patterns such as longevity and strong breeding site

fidelity, providing misleading representation of relative

abundance among species, and sites, and years.

Hydroperiod, water depth, and underwater dynamics of

competition and predation also heavily influence

annual recruitment and population fluctuations (Heyer

1976; Maiorana 1976), and further confound data

interpretation.

As an example, preliminary results of a study comparing

herpetofaunal use of isolated, ephemeral ponds using

intermittent drift fences with pitfall and funnel traps,

indicate significant differences (p < 0.05) in the use of

ponds within regularly burned sandhills (n = 4) versus

hardwood-invaded sandhills (n = 4) by some

amphibian species. Whereas significantly more adult

eastern narrowmouth toads, leopard frogs, and southern

toads used ponds within hardwood-invaded sandhills

during 1994-1997, more adult striped newts used ponds

in savanna-like sandhills (Table 3). Juvenile recruitment

by oak toads and narrowmouth toads was higher in

hardwood-invaded sandhills (Greenberg, unpubl. data),

and recruitment by Florida gopher frogs was higher in

savanna-like sandhills (Greenberg 2001) (Table 3).

However, confidence in the biological significance of

these results is compromised by a lack of clean

distinction between upland matrix treatments. The level

of hardwood invasion in the hardwood-invaded upland

matrix is patchy (Table 1). Further, the distance (on one

side) between 3 ponds in the hardwood-invaded upland

matrix and the savanna-like upland matrix is < 30 m.

Variation in pond use and recruitment among species,

sites, and years is high. These and confounding factors

associated with using ponds as trap foci suggests that

detection of biologically meaningful differences in pond

use among different habitat conditions requires longterm data and heavy replication.

In recent decades, recognition that fire is critical to xeric

pineland ecosystems has led to implementation of

control burning programs for sandhill on many public

lands, where resources permit. In sandhills, the

restoration goal commonly is to create a “park-like”

environment dominated by longleaf pine and wiregrass,

and virtually devoid of a hardwood midstory. This

vision is inspired by some descriptions by early

surveyors and naturalists (e.g., Vignoles 1823; Williams

1837; Harper 1911; Myers 1990), and by the narrow
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 95

Table 3.—Mean (+ SE) adult use (entering and exiting individuals) and juvenile recruitment (exiting individuals

only) of select pond-breeding amphibians using isolated, ephemeral ponds in hardwood-invaded (HI) (n = 4) and

savanna-like (SL) (n=4) longleaf pine-turkey oak-wiregrass sandhills during February 1994 - January 1998, Ocala

National Forest, Florida

 Adults (No./100 trapnights) Juveniles (No./100 trapnights)

 Treatment Treatment

Species Year HI SL P < 0.05 HI SL P < 0.05

Oak Toad 1 0.61 + 0.1 0.12 + 0.08 0.41 + 0.13 0.01 + 0.00 *

2 0.19 + 0.09 0.09 + 0.05 0.00 + 0.00 0.00 + 0.00

3 0.52 + 0.51 0.11 + 0.06 0.02 + 0.02 0.00 + 0.00

4 0.01 + 0.00 0.08 + 0.04 0.00 + 0.00 0.00 + 0.00

5 0.77 + 0.57 0.25 + 0.18 0.12 + 0.12 0.00 + 0.00

Southern Toad 1 0.17 + 0.07 0.03 + 0.01 * 0.00 + 0.00 0.00 + 0.00

2 0.21 + 0.03 0.09 + 0.04 0.00 + 0.00 0.00 + 0.00

3 0.17 + 0.06 0.10 + 0.02 0.15 + 0.14 0.04 + 0.04

4 0.09 + 0.01 0.08 + 0.04 0.00 + 0.00 0.00 + 0.00

5 0.20 + 0.11 0.10 + 0.02 2.82 + 2.79 0.02 + 0.02

Eastern Narrowmouth Toad 1 1.27 + 0.30 0.50 + 0.25 * 0.16 + 0.07 0.01 + 0.01 *

2 0.47 + 0.16 0.27 + 0.09 0.01 + 0.01 0.00 + 0.00

3 0.25 + 0.11 0.19 + 0.07 0.01 + 0.00 0.00 + 0.00

4 0.13 + 0.03 0.18 + 0.09 0.01 + 0.01 0.00 + 0.00

5 0.13 + 0.03 0.07 + 0.04 0.01 + 0.01 0.00 + 0.00

Striped Newt 1 0.00 + 0.00 0.01 + 0.01 * 0.00 + 0.00 0.01 + 0.01

2 0.00 + 0.00 0.00 + 0.00 0.00 + 0.00 0.00 + 0.00

3 0.00 + 0.00 0.00 + 0.00 0.00 + 0.00 0.01 + 0.01

4 0.00 + 0.00 0.10 + 0.08 0.00 + 0.00 0.01 + 0.01

5 0.05 + 0.04 0.26 + 0.23 0.06 + 0.03 1.23 + 1.19

Florida gopher frog 1 0.04 + 0.01 0.02 + 0.01 0.00 + 0.00 0.01 + 0.00 *

2 0.01 + 0.00 0.02 + 0.01 0.15 + 0.04 0.60 + 0.27

3 0.01 + 0.00 0.02 + 0.01 0.07 + 0.03 0.33 + 0.14

4 0.00 + 0.00 0.01 + 0.00 0.01 + 0.01 0.06 + 0.02

5 0.01 + 0.00 0.01 + 0.01 0.32 + 0.04 0.72 + 0.27

Bullfrog 1 0.00 + 0.00 0.01 + 0.00 0.02 + 0.01 0.06 + 0.04

2 0.00 + 0.00 0.00 + 0.00 0.01 + 0.01 0.01 + 0.00

3 0.00 + 0.00 0.00 + 0.00 0.02 + 0.01 0.03 + 0.00

4 0.00 + 0.00 0.00 + 0.00 0.01 + 0.01 0.01 + 0.01

5 0.00 + 0.00 0.00 + 0.00 0.10 + 0.06 0.33 + 0.24

Pig Frog 1 0.00 + 0.00 0.00 + 0.00 0.00 + 0.00 0.00 + 0.00

2 0.00 + 0.00 0.00 + 0.00 0.03 + 0.02 0.04 + 0.02

3 0.01 + 0.00 0.00 + 0.00 0.02 + 0.01 0.07 + 0.03

4 0.00 + 0.00 0.00 + 0.00 0.03 + 0.02 0.15 + 0.11

5 0.00 + 0.00 0.00 + 0.00 0.05 + 0.02 0.03 + 0.02

Leopard Frog 1 0.09 + 0.02 0.07 + 0.04 * 0.25 + 0.13 0.55 + 0.50

2 0.06 + 0.02 0.04 + 0.01 0.11 + 0.03 0.14 + 0.07

3 0.04 + 0.01 0.03 + 0.00 0.12 + 0.04 0.11 + 0.03

4 0.01 + 0.00 0.01 + 0.00 0.01 + 0.01 0.01 + 0.01

5 0.03 + 0.01 0.01 + 0.01 0.40 + 0.10 0.17 + 0.11

Spadfoot Toad 1 0.31 + 0.21 0.73 + 0.47 10.70 + 10.68 0.00 + 0.00

2 0.37 + 0.34 0.08 + 0.07 4.58 + 4.54 0.00 + 0.00

3 0.10 + 0.10 0.02 + 0.01 0.06 + 0.05 0.00 + 0.00

4 0.00 + 0.00 0.00 + 0.00 0.00 + 0.00 0.00 + 0.00

5 0.19 + 0.17 0.10 + 0.05 0.00 + 0.00 0.00 + 0.00
96 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

habitat requirements of the endangered red-cockaded

woodpecker (Carter et al. 1995). However, such

descriptions may over-represent this sandhill variant

because it was more easily (and commonly) traveled

(Landers et al. 1990).

Palynological evidence suggests that the relative

dominance of oak and pine has shifted many times

within the past 20,000 years in the southeastern United

States as a result of climatic and sea level changes that

affects soil moisture and fire frequency (Watts 1971).

Descriptions by early naturalists (see Myers 1990), plant

composition and characteristics, and the presence of

old-growth oaks suggest that even within recent

centuries, fire in sandhills was spatially and temporally

variable (Greenberg and Simons 1999; also see Myers

1990). Prior to fire suppression, patchiness in the

intensity, season, frequency, and spatial extent of fire

allowed some young oaks to reach fire-resistant size and

occur in varying densities over time and across the

sandhills landscape (Greenberg and Simons 1999).

Indeed, several lines of evidence suggest that tree-sized

oaks were an integral part of the sandhill landscape

(Greenberg and Simons 1999). Several oak species,

including bluejack, turkey, and sand post oak are

endemic to xeric pinelands, suggesting that they

historically occurred there. The ability to reach tree size

and reproduce sexually (as well as clonally) indicates

that trees reached maturity frequently enough to retain

the trait (Berg and Hamrick 1994; Landers et al. 1990;

Myers 1990). The presence of acorn-dependent, xericpineland species such as Sherman’s fox squirrels and

red-headed woodpeckers suggests that hard mast was

produced in sandhills. Clearly, stand conditions within

these ecosystems historically were not homogeneous,

but a heterogeneous matrix of stand ages and structural

conditions.

The presence of both specialized, xeric-adapted species

and wide-ranging generalist species in sandhills and

scrub further indicate that historically, variable burning

intensities, intervals, and spatial extent created a range of

microhabitats and stand conditions (Campbell and

Christman 1982). Braithwaite (1987) reported that

lizard species in the wet-dry tropics of Australia were

differentially favored in habitats created by various fire

regimes. He suggested that no single fire regime was

optimal for all species, but that a range of burning

regimes would retain the entire lizard community.

Historically, fire frequency, intensity, and behavior as it

interacted with fuel loads, wind, precipitation,

topography, and pre-existing vegetation patterns was

unlikely to sculpt homogeneous habitat structures

across large landscapes. Whereas the scale of

heterogeneity in stand age, tree or shrub density, and

associated habitat features differed between sandhills

(smaller habitat patches) and scrub (larger habitat

patches), both ecosystems were a mosaic of stand

conditions that shifted spatially and temporally across

the landscape (Greenberg et al. 1994). Populations of

xeric adapted reptiles probably became locally extinct, or

persisted in low numbers within long-unburned habitat

patches, exploiting young, open, suitable habitat

through colonization and reproduction as it was created

by fire.

Ideally, ecosystem management systems should be

designed to incorporate silvicultural and land

management systems that mimic natural disturbance

(Hansen et al., 1991; Greenberg et al., 1994). A

necessary corollary is the need to identify habitat

characteristics that promote diversity and abundance of

species (Hansen et al., 1991). More research on

herpetofaunal response to natural disturbance and the

associated changes in habitat structure is critically

needed in order to gauge the success or failure of

management.

Acknowledgments

I thank C.L. Hardy, L.A. Brennan and their cooperators,

and W. Meshaka and J. Layne for their willingness to

share preliminary or unpublished study results for

mention in this paper. I also thank L.A. Brennan for

presenting this paper at The Wildlife Society conference,

which I was unable to attend. I thank G.W. Tanner, J.

Beach, S.A. and D.A. Johnson, J.S. Staiger, J. Barichivich

the numerous people who have assisted in the

administration, installation, and gathering of field data

for an ongoing study of ephemeral pond use by

amphibians in Florida sandhills. The U.S.D.A. Forest

Service, including the Ocala National Forest, the

Longleaf Pine Ecosystem Restoration Program, and the

Southern Research Station’s Bent Creek Experimental

Forest, and also the Department of Energy and

Savannah River Institute contributed funding to the first

five years of the pond study.

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The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 99

List of Species Names

Vegetation

Wiregrass Aristida stricta (Micheaux)

Turkey oak Quercus laevis (Walt.)

Bluejack oak Quercus incana (Bartr.)

Sand post oak Quercus margaretta (Ashe)

Longleaf pine Pinus palustris (Mill.)

Sand pine Pinus clausa (Chapm. Ex Engelm.) Vasey ex Sarg.

Birds

Red-cockaded woodpecker Picoides borealis

Red-headed woodpecker Melanerpes erythrocephalus

Mammals

Sherman’s fox squirrels Sciurus niger shermani

Herpetofauna

Six-lined racerunner Cnemidophorus sexlineatus

Gopher tortoise Gopherus polyphemus

Short-tailed snake Stilosoma extenuatum

Pine snake Pituophis melanoleucus

Scrub lizard Sceloporus woodi

Sand skink Neoseps reynoldsi

Mole skink Eumeces egregius

Southeastern five-lined skink Eumeces inexpectatus

Ground skink Scincella laterale

Peninsula crown snake Tantilla relicta

Oak toad Bufo quercicus

Southern toad Bufo terrestris

Striped newt Notophthalmus perstriatus

Spadefoot toad Scaphiopus holbrookii

Narrowmouth toad Gastrophryne carolinensis

Florida gopher frog Rana capito aesopus

Pig frog Rana grylio

Bullfrog Rana catesbeiana

Leopard frog Rana utricularia
100 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Abstract.—Use of prescribed fire is increasing in

California oak woodlands, but its effects on vertebrate

wildlife are unknown. We conducted a light-intensity

prescribed fire in mixed blue oak–coast live-oak

woodlands in coastal-central California and assessed

vegetation change and numerical response of small,

nongame vertebrates to the fire. Four of 13 vegetation

and habitat components that we measured were reduced

significantly (P < 0.05) by the fire. We observed no

change in relative abundance of small mammals,

breeding birds, amphibians, or reptiles in response to

the prescribed fire.

Introduction

Oak (Quercus spp.) woodlands are the most extensive

vegetation type in California, covering approximately

three million ha (Rossi 1979; Griffin and Muick 1984).

In varying compositions, 10 native shrub and nine

native tree species comprise numerous woodland

habitat types. More vertebrate wildlife species use oak

woodlands than any other vegetation type in California

(Ohmann and Mayer 1987; Airola 1988).

California oak woodlands are fire-adapted, having

evolved with fire during the past one million years. The

dominant use of oak woodlands is rangeland for

livestock production. Prescribed fire is used in oak

woodlands as a livestock forage and fuel management

tool (Griffin and Muick 1984). The California Fire Plan

(California Department of Forestry and Fire Protection

1996) suggests increased use of prescribed fire in

California’s oak woodlands to reduce the severity of

wildfires by limiting fuel accumulation and to manage

livestock forage.

In the central coast region of California (roughly the

area between Santa Barbara and San Francisco extending

to the coast mountain ranges approximately 80 km

inland), the California Department of Forestry and Fire

Protection (CDF) conducts prescribed burns on 1,000–

4,000 ha annually (Ben Parker, CDF, San Luis Obispo

Ranger Unit, San Luis Obispo, CA, pers. comm.). The

CDF presumes that prescribed fire benefits wildlife of

oak woodland habitats, but no published research

supports this assumption. Most research on effects of

prescribed fire to wildlife and their habitats in California

historically has been conducted in chaparral ecosystems

(Lawrence 1966; Lillywhite 1974; Longhurst 1978;

Quinn 1979, 1983; Wirtz 1979, 1982). Except for a

manuscript from this project (Vreeland and Tietje

1998), we are aware of no published research on effects

of prescribed fire to California oak woodland habitats or

associated wildlife species. Our objective was to quantify

vegetation change and numerical response of small,

nongame vertebrates to a prescribed fire conducted in

mixed oak woodlands of the central coast region of

California.

Study Area

We conducted this study at Camp Roberts (CR),

California, a facility of the California Army National

Guard located approximately 30 km from the Pacific

Ocean in northern San Luis Obispo and southern

Monterey counties. Topography at CR varies from flat to

gently rolling hills and steep (>45° slope) hills. The

climate of the study area is Mediterranean, characterized

by cool, wet winters and hot, dry summers. Annual

precipitation averages 38 cm (66-year range=10.8–74.1

cm), falling almost exclusively as rain between

November and March. Mean annual temperature

averages 15.3°C. Summer high temperatures frequently

approach 50°C; winter lows rarely fall below –6°C.

Sheep grazing and military training occur in our study

area, but are limited in extent, duration, and intensity.

Fire has been excluded from the study area for >15 years

(Brian Duke, CR Environmental Office, Camp Roberts,

CA). Our study sites were on slopes <20°, on north- to

east-facing aspects, and 300–500 m elevation.

Camp Roberts covers 17,000 ha, with 41% classified as

oak woodland (Camp Roberts EMAP 1989). Three oak

habitat types occur at CR: valley oak (Q. lobata), coast

live-oak (Q. agrifolia), and blue oak (Q. douglasii), with

considerable overlap between coast live-oak and blue

oak types.

We used blue oak and mixed blue oak–coast live oak

stands in the San Luis Obispo County portion of CR

(Tietje et al. 1997; Tietje and Vreeland 1997a). Blue oak

was the dominant canopy species with a variable

contribution (0–45%) of coast live-oak. Where present,

understory vegetation was composed primarily of toyon

(Heteromeles arbutifolia), redberry (Rhamnus crocea),

poison oak (Toxicodendron diversilobum), and bigberry

manzanita (Arctoctsphylos glauca). Small (≤ 0.25 ha)

patches of chaparral (Ceonothus spp. and Adenostoma

fasciculatum) occurred on three study plots. Common

forbs included hummingbird sage (Salvia spathacea),

deerweed (Lotus scoparius), filaree (Erodium spp.), and

fiddleneck (Amsinckia spp.). Wild oats (Avena spp.),

Numerical Response of Small Vertebrates to Prescribed Fire

in California Oak Woodland

Justin K. Vreeland1 and William D. Tietje2

1

Pennsylvania Cooperative Fish and Wildlife Research Unit,

University Park, PA 16801

2Department of Environmental Science, Policy, and

Management, University of California, Berkeley, CA 94720
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 101

bromes (Bromus spp.), and fescues (Festuca spp.)

dominated grassy openings of the woodland floor.

Blue oak sites were characterized by 40–60% canopy

cover, <10% shrub cover, and >50% ground cover of

exotic annual grasses. Mixed blue oak–coast live-oak

sites were characterized by 60–90% canopy cover, 10–

30% shrub cover, and a well-developed, thick leaf litter

layer with abundant herbs, forbs, and <30% grass cover.

Tietje et al. (1997) and Tietje and Vreeland (1997a) give

detailed descriptions of vegetative and habitat

characteristics of our study sites.

Methods

Prescribed Fire

We selected two areas to receive a prescribed fire

treatment. One area (hereafter “Burn 1”) was 80 ha, the

other area (hereafter “Burn 2”) was 120 ha. The

California Department of Forestry and Fire Protection

burned these areas on consecutive days in October

1997. The perimeter was burned first, then the interiors

were ignited using drip torches, and accelerants

delivered from helicopters. Surrounding unburned areas

hereafter are referred to as “Ctrl” or “unburned areas.”

Vegetation Sampling

We sampled vegetation in 11 1-ha plots throughout the

burned areas within two months before and within 2–3

months after the prescribed fire. We randomly located

five 10-m-radius sampling stations on each plot (Figure

1). At 10 m in the cardinal directions from the central

point of each station, we measured canopy cover with a

concave spherical densiometer (Lemmon 1956),

understory obstruction (an index to shrub cover) with a

vegetation pole (Griffith and Youtie 1988) graduated in

five 0.5-m sections, and ocularly estimated ground cover

within 1-m2 frames. Within 10 m of the central point of

each station, we counted the number of pieces of coarse

woody debris (CWD, 31 m long and maximum diameter

310 cm), number of snags (31.37 m tall and 310 cm

diameter at breast height), and number of dwellings

(hereafter “houses”) constructed by dusky-footed

woodrats (Neotoma fuscipes).

Small Mammal Sampling

We live-trapped small mammals on 22 1-ha grids (8 x 8

grids, 15-m spacing) during May, 1997–1999. Plots were

evenly divided between burned and unburned areas; six

plots were in Burn 1 and five plots were in Burn 2. Plots

were 342 m apart. Spatial separation between plots was

adequate because <4 animals moved among plots

during the course of the study (W.D. Tietje, unpublished

data). We trapped for five consecutive nights during

1997 and three consecutive nights during 1998 and

1999. We baited traps with horse feed (“COB”: corn,

oats, and barley laced with molasses), tagged animals

with individually numbered ear tags, and released them

Ground

Canopy 

Shrub

CWD

Snags

Woodrat 

House

N

S

W E

10 m

Figure 1.—Schematic layout of 10-m, circular plots used to sample vegetation characteristics

in blue oak–coast live oak woodlands in San Luis Obispo County, California, 1997–1998.

“Ground” represents ocular estimation of ground cover in 1 m2

 quadrats. “Canopy”

represents measurement of canopy cover by concave spherical densiometer (Lemmon 1956).

“Shrub” represents measurement of understory obstruction or shrub cover with a vegetation

sampling pole (Griffith and Youtie 1988). “CWD,” “Snags,” and “Woodrat House” represent

counting these elements within 10 m of plot centers. Not drawn to scale.
102 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

at site of capture. We handled animals in accordance

with University of California, Berkeley Animal Use

Protocol #R166-0199.

Breeding Bird Sampling

We conducted point-counts of breeding birds during

March–April, 1997–1999. We used 86 50-m-radius

point count stations located 3150 m apart. Stations were

evenly divided between burned (13 stations were in

Burn 1, 30 stations were in Burn 2) and unburned areas.

We visited each station six times in each season. We

rotated station visitations among start time, observer,

and treatment. Two to four observers conducted counts

in each year. We conducted counts for 10 minutes at

each station between official sunrise and 1100 hours

using standard breeding-bird survey protocols (Bibby et

al. 1992).

Amphibian and Reptile Sampling

We counted amphibians and reptiles under 136 61-cm x

61-cm x 1.27-cm plywood coverboards (DeGraaf and

Yamasaki 1992; Grant et al. 1992; Tietje and Vreeland

1997b) on each of nine 5.8-ha plots (8 x 17 grid, 30- x

15-m spacing). Five plots were in the unburned area,

two plots each were in Burn 1 and Burn 2. We checked

each coverboard once every 7–10 days during late

January–early May in 1995–1999. We first deployed

coverboards in 1994 to allow them to weather for 6–9

months, dissipating chemicals used in their manufacture

that might affect their use by amphibians and reptiles

(Grant et al. 1992). We had difficulty identifying slender

salamanders (Batrachoseps spp.) (N. Scott, U.S.

Geological Survey, Biological Resources Division, San

Simeon, CA, pers. comm.). Pacific slender salamanders

(B. pacificus) and black-bellied salamanders (B.

nigriventris) potentially occur at CR, but cannot be

distinguished in the field. Therefore we grouped them

into one species category, hereafter “slender

salamanders.”

Analyses

We qualitatively described fire behavior and counted the

number of grid intersections on small mammal trapping

grids where fire burned to within 1 m to quantify areal

coverage of the fire. In Vreeland and Tietje (1998), we

conducted paired-sample t-tests to assess vegetation

changes and considered differences significant at

α=0.05. We repeat an abbreviated version of those

vegetation results here, but direct the reader to Vreeland

and Tietje (1998) for details on vegetation results.

Because we were unable to randomly select burned areas

and because treatment replication was low, our animal

data cannot meet some assumptions of analyses of

variance (ANOVA). Accordingly, rather than conduct

inappropriate, low-power statistical tests, we used means

and 95% confidence intervals (CI) to assess numerical

response of vertebrates to the prescribed fire. We selected

one species to represent each of the four taxa we

monitored. We selected a representative species based on

two characteristics: large relative abundance and

potential for demonstrating change as a result of

prescribed fire. True replication in our study is at the

burn unit level. Relative abundance was calculated for

each sampling unit (plot or point count station), then

averaged for a treatment area (Burn 1, Burn 2, Ctrl).

Because we have pre- and post-fire data on burned and

unburned areas, pre-fire differences in relative

abundance between burned and unburned areas are

unimportant. The meaningful test of an effect of the fire

is a comparison between relative abundance of a species

on burned areas compared to unburned areas before

and after the fire. This is analogous to the interaction

term in a 2-way ANOVA. Differences between treatment

means were considered significant if 95% of the CI did

not overlap.

Results

Prescribed Fire

The California Department of Forestry and Fire

Protection rated the prescribed fire as light- to moderateintensity (3–4 on a 10-point scale). Flame height

generally was <1 m except in a few areas of decadent

grasses, chaparral, or dense accumulations of CWD. Few

mature trees died. The fire was patchily distributed and

carried better through blue oak stands with abundant

grass cover than through mixed oak stands with thick

leaf litter and dense canopy cover. Mineral soil was

mostly unexposed except in small areas with high fuel

load (dense CWD, chaparral). Area burned on 1-ha

plots averaged 46% and ranged from 30 to 66%.

Vegetation

Understory obstruction and grass cover were reduced by

7% and 70%, respectively. Much of the reduction in

understory obstruction was the result of reduction in

grass cover that covered the lower 1 m of the vegetation

pole. Excluding this grass cover probably would result in

no statistical reduction in understory obstruction. Grass

cover returned to pre-fire cover by one growing season

after the fire. Canopy cover, number of snags, and leaf

litter depth (an index to litter volume), did not change.

Approximately six % of the canopy was singed. Number

of woodrat houses was reduced by 30%. Number of

pieces of CWD was reduced by 35%. We recorded five

new pieces of CWD in the 55 10-m vegetation plots.

These pieces of CWD were created when roots of mature

oaks were burned during the fire, causing the bole to fall

to the ground. Vreeland and Tietje (1998) report

detailed vegetation responses to the prescribed fire.

Small Mammals

We captured nine species of small mammals during

three years of live-trapping (Table 1). Woodrats, piñon
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 103

Table 1.—Relative abundance of small mammals, breeding birds with at

least 100 detections, amphibians, and reptiles monitored in response to

prescribed fire in blue oak–coast live oak woodlands in San Luis Obispo

County, California, 1995–1999

Common name (scientific name) Relative abundancea

Small Mammals

Dusky-footed woodrat (Neotoma fuscipes) 26.49

Piñon mouse (Peromyscus truei) 6.61

Brush mouse (Peromyscus boylii) 4.40

California mouse (Peromyscus californicus) 3.06

California pocket mouse (Perognathus californicus) 1.34

California vole (Microtus californicus) 0.36

Merriam’s chipmunk (Tamias merriami) 0.36

Deer mouse (Peromyscus maniculatus) 0.11

Heermann’s kangaroo rat (Dipodomys heermanni) 0.01

Breeding Birdsb

Oak titmouse (Parus inornatus) 3,001

Dark-eyed junco (Junco hyemalis) 2,564

Lesser goldfinch (Carduelis psaltria) 1,842

Bushtit (Psaltriparus minimus) 1,529

Western scrub-jay (Aphelocoma coerulescens) 1,344

House finch (Carpodacus mexicanus) 1,336

Bewick’s wren (Thryomanes bewickii) 1,221

Violet-green swallow (Tachycineta thalassina) 805

Spotted towhee (Pipilo erythrophthalmus) 643

Anna’s hummingbird (Calypte anna) 517

California quail (Callipepla californica) 505

White-breasted nuthatch (Sitta carolinensis) 485

Hutton’s vireo (Vireo huttoni) 480

Yellow-rumped warbler (Dendroica coronota) 480

Golden-crowned sparrow (Zonotrichia atricapilla) 459

Ash-throated flycatcher (Myiarchus cinerascens) 453

California towhee (Pipilo crissalis) 426

Western bluebird (Sialia mexicana) 405

Nuttall’s woodpecker (Picoides nutallii) 394

Mourning dove (Zenaida macroura) 350

Ruby-crowned kinglet (Regulus calendula) 315

Blue-gray gnatcatcher (Polioptila caerulea) 285

Acorn woodpecker (Melanerpes formicivorus) 246

Orange-crowned warbler (Vermivora celata) 245

Wrentit (Chamaea fasciata) 204

Lawrence’s goldfinch (Carduelis lawrencei) 177

Hermit thrush (Catharus guttatus) 141

Lark sparrow (Chondestes grammacus) 134

Northern flicker (Colaptes auratus) 121

Brown-headed cowbird (Molothrus ater) 111

Amphibians and Reptiles

Western skink (Eumeces skiltonianus) 3.96

Western fence lizard (Sceloporus occidentalis) 2.19

Slender salamander (Batrachoseps sp.) 1.74

California legless lizard (Anniella pulchra) 0.47

continued
104 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

mice (Peromyscus truei), brush mice (P. boylii), and

California mice (P. californicus) were the four most

abundant species. Woodrats were the most frequently

captured species (26.5 captures/100 trap-nights during

1997–1999). Relative abundance (captures/100 trapnights) of woodrats on burned areas compared to the

unburned areas did not change after the prescribed fire

(Figure 2, Table 2). Relative abundance of woodrats was

approximately 60% lower in 1998 and 1999 than in

1997, but this reduction was proportional among

burned and unburned areas. Confidence intervals

consistently overlapped treatment means and other CI

within and among years.

Breeding Birds

We observed over 85 species of birds during the

breeding season on all 86 point count stations (Table 1).

Dark-eyed juncos (Junco hyemalis) were one of the two

most frequently observed species (1.7 observations/

point/visit during 1997–1999, respectively). Relative

abundance (mean number of observations/point) of

juncos on burned areas compared to unburned areas did

not change after the prescribed fire (Figure 3, Table 2).

We observed an approximately 80 % increase in juncos

two years post-fire, but this increase was proportional

among burned and unburned areas. Juncos consistently

were more abundant on Burn 1 than Ctrl, but the

difference remained consistent before and after the fire.

Other confidence intervals were narrow, but consistently

overlapped treatment means and other CI within and

among years.

Amphibians and Reptiles

We observed four amphibian (two salamanders, one

frog, one toad) and 11 reptile (five lizards, six snakes)

species under plywood coverboards (Table 1). Among

amphibians, only slender salamanders were observed in

appreciable numbers (1.74 observations/100 coverboard

visits during 1995–1999). Relative abundance (number

of observations/100 board-visits) of slender salamanders

was similar on burned areas compared to unburned

areas before and after the prescribed fire (Table 2).

Except during 1999, slender salamanders were more

abundant on Burn 1 than Burn 2 or Ctrl, but this

pattern of relative abundance remained consistent

among years before and after the prescribed fire.

Confidence intervals were broad, frequently included

zero and other treatment means, and overlapped other

CI within and among years.

Western skinks (Eumeces skiltonianus) were the most

frequently observed reptile species (3.96 observations/

100 coverboard visits during 1995–1999). Relative

abundance (number of observations/100 board-visits)

of western skinks varied among years and among

treatment areas within and among years, but was similar

on burned areas compared to unburned areas before

and after the prescribed fire (Table 2). Skink observation

rate appeared to be increasing on Ctrl, decreasing on

Burn 2, and irregular on Burn 1, but these trends began

2–3 years before the fire, and were not caused by nor

interrupted by the fire. Confidence intervals were broad,

frequently included zero and other treatment means,

and overlapped other CI within and among years.

Southern alligator lizard (Gerrhonotus multicarinatus) 0.35

Gopher snake (Pituophis melanoleucus) 0.22

California whipsnake (Masticophis lateralis) 0.22

Ring-necked snake (Diadophis punctatus) 0.14

Night snake (Hypsiglena torquata) 0.07

Common king snake (Lampropeltis getulus) 0.06

Chorus frog (Hyla regilla) 0.06

Side-blotched lizard (Uta stansburiana) 0.03

Western toad (Bufo boreas) 0.02

Common garter snake (Thamnophis sirtalis) 0.01

Monterey salamander (Ensatina eschscholtzii) 0.01

a

Small mammals: captures/100 trap-nights; breeding birds: total observations

during 3 years (86 total point count stations); amphibians and reptiles:

observations/100 coverboard checks.

b

Fifty-five species of birds were observed fewer than 100 times during 3 years

and are not shown.

Table 1.—continued

Common name (scientific name) Relative abundancea
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 105

Discussion

Conclusions

We detected no change in relative abundance of small

mammals, breeding birds, amphibians, or reptiles after a

light- to moderate-intensity prescribed fire in California

blue oak–coast live oak woodlands. Intensity of the fire

was limited by cool temperatures, moderately high

relative humidity, low wind speed, and resulted in

generally marginal or short-term changes to the

vegetation characteristics we measured. Prescribed fire in

oak woodlands may reduce resource competition from

exotic annual grasses, stimulating shrub and tree health

and vigor and, ultimately, mast production and overall

habitat rejuvenation. Small vertebrates may respond to

this habitat rejuvenation.

Coarse woody debris serves as hiding, breeding, and

foraging habitat for small vertebrates. Reduction in

CWD was substantial after the prescribed fire, but CWD

may not be limiting for animals that use it. We counted

237 pieces of CWD on 1.73 ha of vegetation sampling

plots (137 pieces/ha). Size criteria for CWD was

appropriate to gauge use by most vertebrates but is a

conservative estimate of CWD abundance, because small

vertebrates will use smaller pieces of CWD. We observed

slender salamanders using pieces of woody debris (e.g.,

dead coast live-oak bark) as small 20 cm x 20 cm x 3 cm,

considerably smaller than the minimum size of CWD

for this study.

Our casual observations detected only one instance (a

woodrat) of direct mortality caused by the fire. Direct

mortality likely was low, because most animals we

monitored have subterranean access or are capable of

other strategies that enable them to escape fire. Russell

et al. (1999) reviewed over 12 published papers on

effects of fire to herpetiles and concluded that direct

mortality of these animals in fires is low.

No other published literature reports on the effects of

prescribed fire to vegetation structure or vertebrate

communities in California oak woodlands. Many

researchers have compared burned and unburned areas

after fire, but lack of pre-fire observations limits

conclusions that can be drawn about effects of fire to

animals. Some studies have combined other silvicultural

treatments (e.g., thinning, forest harvesting) with

burning. Distinguishing effects of burning from other

treatments can be difficult, especially if treatments were

not conducted under mutually exclusive study designs.

Much study of prescribed fire and its effects on habitat

Figure 2.—Dusky-footed woodrat abundance

(captures per 100 trap-nights) in May 1997–1999

before and after an October, 1997 prescribed fire

in blue oak–coast live oak woodlands in San Luis

Obispo County, California. Error bars are 95%

confidence intervals.

0

20

40

60

80

1997 1998 1999

Caps / 100 T-N

BURN1 (n=6)

BURN2 (n=5)

CTRL (n=11)

Figure 3.—Dark-eyed junco abundance (mean

number of observations per point) in March and

April, 1997–1999 before and after an October,

1997 prescribed fire in blue oak–coast live oak

woodlands in San Luis Obispo County, California.

Error bars are 95% confidence intervals.

0

5

10

15

20

25

30

35

40

1997 1998 1999

BURN1 (n=13)

BURN2 (n=30)

CTRL (n=43)

Mean Obs. / Point
106 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

structure and wildlife have been conducted in other

ecosystems (e.g., California chaparral, California dry

pine, southeastern U.S. pine forests, prairie, northern

temperate oak savannah, Appalachian hardwoods, and

others). Species in different habitats and ecosystems

likely respond differently to prescribed fire; therefore,

comparisons of our results to other prescribed fire

studies in other habitat types are hampered by these

cross-habitat differences. Therefore, we restrict our

comparisons with other prescribed fire studies to those

that experimentally investigated effects of fire, rather

than with studies that conducted post-fire observations

on burned and unburned areas, on burned areas before

and after fire without unburned controls, and wildfires

that generally burn hotter and faster than prescribed fires.

Despite a 30% reduction in number of woodrat houses,

we observed no change in relative abundance of

woodrats because of the prescribed fire. Not all woodrat

houses are occupied (Vreeland and Tietje 1999) and fire

did not reach many houses in the densest habitats;

therefore, few woodrats were forced to relocate to new

areas or construct new houses after the fire. Tevis (1956)

experimentally investigated effects of a site-preparation

fire in logging debris (Pacific madrone [Arbutus

menziesii] and tanoak [Lithocarpus densiflorus]) on

numbers of small mammals in northern California

Douglas-fir (Pseudotseuga menziesii) stands. Over 70 % of

marked animals were not recaptured after the fire and

probably died or emigrated from the burned area

immediately after the fire. Most recaptured animals were

caught at the periphery of the fire, suggesting they were

not resident within the burn blocks before the fire or

temporarily relocated after the fire. However, within

three weeks after the fire, more new individuals were

captured in the burned area than were captured before

the fire. Tevis (1956) attributed the immigration by new

individuals to the lack of resident mice created by the

fire. We observed no such immigration to burned areas,

probably because few or no residents were killed by the

fire or emigrated from the burned areas, thereby leaving

little unoccupied habitat.

Similarly, Tester (1965) trapped deer mice (P.

maniculatus and P. leucopus) before and after a prescribed

fire in burr oak (Q. macrocarpa)–northern pin oak (Q.

ellipsoidalis) savanna in Minnesota and reported

substantial immigration to burned areas within one

month after the fire. Few animals captured before the

fire were recaptured after the fire, but Tester (1965)

could not speculate if absence of these animals

suggested low survival during the fire, emigration, or

high predation following the fire. Tester (1965)

attributed increased use of burned areas to reduction in

litter cover and depth, which were negatively correlated

with presence of deer mice.

Table 2.—Relative abundance (mean, 95% CI) of dusky-footed woodrats, dark-eyed juncos, slender

salamanders, and western skinks on burned and unburned areas before and after prescribed fire in

blue oak–coast live oak woodlands in San Luis Obispo County, California, 1995–1999. The

prescribed fire was conducted in October, 1997. Pre-fire years are 1995–1997 for salamanders and

skinks, and 1997 for woodrats and juncos. Woodrats and juncos were not monitored in 1995 and

1996. See text for sample sizes

 Year Treatment Dusky-footed Dark-eyed Slender Western

woodrat junco salamander skink

1995 Burn 1 8.6 (0–89.0) 2.6 (00.–13.3)

Burn 2 0.8 (0–07.7) 4.0 (00.–08.3)

Ctrl 0.6 (0–01.4) 3.3 (00.–06.7)

1996 Burn 1 2.0 (0–15.1) 2.0 (00.–12.3)

Burn 2 1.0 (0–11.3) 3.4 (00.–10.4)

Ctrl 0.3 (0–00.6) 3.5 (00.–07.0)

1997 Burn 1 25.8 (13.2–38.3) 15.6 (11.3–19.9) 2.9 (0–19.2) 1.7 (00.–07.7)

Burn 2 47.6 (36.1–59.1) 08.4 (06.9–09.8) 2.4 (0–21.6) 2.7 (00.–09.7)

Ctrl 31.2 (18.8–43.5) 07.4 (06.0–08.9) 1.0 (0–02.2) 4.8 (0.2–09.3)

1998 Burn 1 10.6 (07.0–14.2) 12.4 (10.1–14.6) 6.6 (0–52.4) 3.3 (00.–26.1)

Burn 2 28.3 (13.6–43.0) 07.8 (06.0–09.6) 3.2 (0–32.1) 2.4 (00.–04.7)

Ctrl 22.1 (10.6–33.6) 06.5 (05.4–07.6) 1.0 (0–02.2) 6.7 (0.3–13.0)

1999 Burn 1 10.3 (00.0–20.7) 26.3 (18.9–33.7) 1.8 (0–15.4) 2.8 (00.–18.9)

Burn 2 27.2 (12.4–42.0) 15.9 (11.8–18.6) 2.8 (0–33.1) 2.4 (00.–15.6)

Ctrl 24.0 (15.4–32.7) 11.6 (08.0–15.2) 0.4 (0–00.9) 6.7 (00.–12.7)
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 107

Masters et al. (1998) observed greater small mammal

abundance, species richness, and diversity in Arkansas

shortleaf pine (Pinus echinata) stands treated with

prescribed fire than in unburned stands. Sullivan and

Boateng (1996) observed dramatic decreases in deer

mice and Oregon voles (Microtus oregoni) immediately

following prescribed fire in coniferous forests of British

Columbia, but animal abundance returned to pre-fire

levels within four months following the fire. Red-backed

voles (Clethrionomys gapperi) also increased substantially

immediately following fire. Kaufman et al. (1988)

observed a doubling of deer mouse abundance on

burned areas 3–5 weeks following prescribed fire in

Kansas tall-grass prairie. Ford et al. (1999) observed a

decrease in abundance of pine voles (Microtus pinetorum)

following prescribed burning in pitch pine (Pinus rigida)

and oak stands in North Carolina, but abundance of

nine other species did not change following fire.

Lawrence (1966) trapped dusky-footed woodrats and

observed immediate, but marginal short-term reductions

in abundance following prescribed fire in chaparral and

grasslands in California. Pattern of abundance between

burned and unburned areas in both vegetation types

remained similar for nearly three years following fire.

Lawrence (1966) also observed decreased mass of piñon

mice on burned areas following fire, but seasonal

pattern of mass was similar between burned and

unburned areas. Lawrence (1966) attributed decreased

mass to 90% reduction in food abundance. Masters et

al. (1998) and Kaufman et al. (1988) attributed

increases in abundance to more favorable habitat or

foraging conditions in burned areas.

Effects of prescribed fires conducted during fall may be

less extensive or severe for breeding bird communities

than for wintering bird communities in California oak

woodlands, because breeding activity and behavior may

commence during or after habitat rejuvenation initiates.

Wilson et al. (1995) observed varied responses by

breeding birds to prescribed fire in shortleaf pine–oak

forests in Arkansas. Overall, bird densities and density of

ground- and shrub-foraging and shrub-nesting species

were greater in burned than in unburned stands

following fire. Similarly, Salveter et al. (1996) observed

no changes in abundance of neotropical migrant or

resident songbird species following experimental

prescribed fire in shortleaf pine and oak forests in

Arkansas. Lawrence (1966) observed immediate

reductions in abundance of chaparral birds following

prescribed fire in California chaparral. Lower bird

abundance persisted in burned areas compared to

unburned areas through three years post-fire.

Conversely, Lawrence (1966) observed increases in

grassland birds and no significant change in abundance

of oak woodland birds during the same fire in chaparral.

Horton and Mannan (1988) observed a reduction in

abundance of northern flickers (Colaptes auratus) and

violet-green swallows (Tachycineta thalassina), an increase

in mountain chickadees (Parus gambeli), and no change

in 13 other species following a low-intensity prescribed

fire in Arizona ponderosa pine (Pinus ponderosa) forests.

We also monitored violet-green swallows and flickers in

our oak-woodland study site, but observed no changes

in their relative abundance, although detection rates

were low. Salveter et al. (1996) suggested that some

guilds (e.g., ground foragers, cavity nesters) may

collectively demonstrate increases or decreases in

abundance following prescribed fire. Reynolds and

Krausman (1998) observed no reduction in relative

abundance of breeding birds or wintering birds

following experimental winter prescribed burning in

Texas mesquite (Prosopis glandulosa) grasslands, and

argued that species respond differently, potentially

confounding guild analyses. Petersen and Best (1987)

observed no immediate reduction in bird density

following prescribed fire in sagebrush (Artemisia spp.)

rangelands in Idaho, but documented increased species

richness and density two years post-fire. Winter and Best

(1985) observed a change in placement position of sage

sparrow (Amphispiza belli) nests following prescribed

burning in sagebrush (Artimesia spp.) rangelands in

Idaho, suggesting functional responses to prescribed fire

may be as important as changes in bird abundance.

Slender salamanders require moist microhabitats to

maintain suitable respiratory conditions, and therefore

are closely associated with habitat components (e.g.,

dense canopy cover, presence of CWD) conducive to

maintaining these conditions. Western skinks are habitat

generalists and ubiquitous, although they likely require

CWD as refugia (Tietje et al. 1997; Tietje and Vreeland

1997a). Because CWD was abundant on our study plots,

CWD probably is not limiting populations of slender

salamanders, western skinks, or other herpetiles.

Therefore, even large reductions in CWD may not result

in detectable changes in relative abundance of small

vertebrates.

We were unable to find experimental studies on effects

of prescribed fire (without other silvicultural

treatments) to amphibians and reptiles. Anecdotal and

observational studies suggest that amphibians and

reptiles may respond negatively (Enge and Marion 1986,

McLeod and Gates 1998) or not at all (Enge and Marion

1986, Ford et al. 1999) to prescribed fire. Other

observational studies suggest reptiles may exhibit

different functional or behavioral activity in burned

areas (Kahn 1960, Lillywhite and North 1974, Lillywhite

et al. 1977). Observation rates of amphibians and

reptiles typically are low in most studies, contributing to

low measurement precision in response to treatments.

Because we did not mark amphibians and reptiles, we

relied on observation rates to quantify their relative

abundance. If herpetile numbers were reduced by the

fire, these reductions could be masked by increased

movement and subsequent increased observation rates,

or increased use of coverboards with fire-induced loss of

natural cover objects. Increased movement rates could

result from changes in prey (e.g., small mammals,
108 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

invertebrates) abundance, foraging behavior, increased

ease of mobility through ground vegetation reduced by

fire, or genuine increases in animal abundance.

Caveats

Replication in our study was low because plots and

point-count stations are pseudoreplicates of the burned

(n=2) and unburned (n=1) areas. Therefore, true

replication in our study was the burn blocks. Replicating

prescribed fire studies at this scale is difficult for

logistical reasons and budget constraints, but

researchers should strive to replicate at the landscape

level.

Sampling units were few for amphibian and reptile data

(four burned plots, five unburned plots), contributing to

low statistical power (broad CI) for these data.

Furthermore, we did not mark individual birds,

amphibians, or reptiles; therefore, our measures of

relative abundance lack precision and may overestimate

relative abundance. Although we individually marked

small mammals, budget constraints restricted us to

trapping only three nights during the two post-fire

trapping sessions, which forced us to use capture rates

rather than number of individuals as the metric for

small mammal relative abundance. Data collected on

number of individuals rather than capture rates has

lower explanatory power.

Research on breeding birds suggests that bird density

and relative abundance are poor measures of habitat

quality and indicators for responses to treatments (Van

Horne 1983, Vickery et al. 1992a,b). Engstrom et al.

(1996) suggest that estimates of productivity and site

fidelity would be better variables than density or relative

abundance for demonstrating potential effects of

prescribed fire to avian communities because relative

abundance or density can be an inappropriate index to

habitat quality.

Despite these shortcomings to our study, we believe our

research represents important, initial information on

effects of prescribed fire to blue oak–coast live-oak

woodland habitat and associated small, nongame

vertebrates. In addition, consistent responses to the

prescribed fire by species in the four taxa we monitored

suggest that our conclusions are appropriate.

We conducted the prescribed fire in October 1997, but

assessed responses of small vertebrates 3–7 months after

the fire. Monitoring animals immediately after the fire

may have resulted in different effects to our study taxa.

Our observations on the response to prescribed fire by

small mammals during fall (October and November)

and by wintering birds (January and February) followed

similar patterns to the monitoring efforts we report here,

however (Vreeland and Tietje 1998; W.D. Tietje, unpubl.

data).

Research Implications

Studies of effects of prescribed fire must maintain an

experimental design: before and after sampling on

burned and unburned areas is required to document

changes in animal numbers on burned areas compared

to unburned controls (Russell et al. 1999). Replicating at

landscape-level scales (>100 ha) is critical to assessing

effects of prescribed fire, which necessarily occurs over

large areas. Effects of prescribed fire, particularly for

more intense fires, may last beyond two years post fire;

long-term studies should evaluate effects of prescribed

fire >5–20 years post-fire. Response of vegetation and

animals may differ among fire intensity and among

seasons in which prescribed fires are conducted

(Engstrom et al. 1996, King et al. 1998). Further study

of effect of season and intensity of fire is warranted.

Researchers should mark individuals, preferably with

individual marks, to precisely measure changes in

numbers of individuals after prescribed fires. In addition

to numerical responses, animals may demonstrate

functional and behavioral responses to prescribed fire

(Kahn 1960, Lillywhite and North 1974, Lillywhite et al.

1977). Researchers should assess functional response of

animals and changes in behavior (foraging and nesting

behavior, movement patterns, home range size) as well

as numerical response (Winter and Best 1985, Engstrom

et al. 1996). Estimates of reproductive potential (nest

success [Engstrom et al. 1996], small mammal testes

size), or individual status (health index [Lawrence

1966]) before and after prescribed fire would greatly

enhance knowledge of effects of prescribed fire to small

vertebrates (Engstrom et al. 1996).

Acknowledgements

We thank CDF (Grant #8CA96037) and the Integrated

Hardwood Range Management Program at University of

California Cooperative Extension (Program Grant #91-

003) for funding this study. Camp Roberts permitted

access to study sites. The California Department of

Forestry and Fire Protection and the Camp Roberts fire

department conducted the prescribed burns. The San

Luis Obispo office of the University of California

Cooperative Extension and Camp Roberts

Environmental Office provided logistical support. We

gratefully acknowledge the work of the many field

biologists that assisted with data collection.

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The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 111

Abstract.—The Nature Conservancy has conducted

prescribed burns for two decades in North Carolina’s

Green Swamp to enhance and maintain species

biodiversity in the longleaf pine (Pinus palustris)

savannas and ecotones. The surrounding pocosins,

however, have been left alone. Increasingly, dense shrub

growth in the pocosin suppresses herbaceous species

and increases the risk of serious wildfires. Two

experimental burns have shown that it is possible to

safely conduct prescribed burns in pocosins. Continuing

studies will monitor the vegetation response to fire and

map the vegetation and fuels to predict fire behavior,

leading to a better fire management plan for the Green

Swamp based on its historic fire regime.

In managing more than 1,500 natural areas (6.8 million

acres) throughout the United States, The Nature

Conservancy (TNC) examines the processes that have

shaped and are shaping the ecosystems targeted for

conservation within its program. The Nature

Conservancy uses prescribed fires where this tool will

contribute to the perpetuation of species and

ecosystems. The Nature Conservancy has increased its

ecological burning for ecological reasons to just over

70,000 acres, or 450 burns per year, in 44 states over the

last decade. Fire exclusion and altered fire regimes have

been identified as threats in 45% of sites where TNC

actively is working and over 600,000 acres presently are

targeted for burning. The North Carolina TNC Chapter

has 14 preserves with 25,000 acres under fire

management, ranging from longleaf and pond pine

(Pinus serotina) ecosystems to Carolina bays, Piedmont

prairie remnants, and a lone mountain seepage bog. In

the 1980’s, TNC began its fire management by burning

longleaf pine savannas during the winter months. This is

the traditional prescribed burning season in the region

because during this time fire behavior is most

predictable and controllable. As the chapter’s staff

gained experience with prescribed burning, a shift to

conducting burns designed and timed to mimic fire

regimes under which the natural communities evolved

occured, also while protecting and promoting associated

indigenous rare species. In North Carolina, this meant

that the TNC’s statewide prescribed burns are now

conducted year-round, but with emphasis on the growing

season to mimic wildfires as a key ecological process.

Role of Fire in the Ecology of

the Green Swamp

The Green Swamp is a 17,000 acre preserve in the

southeastern Coastal Plain of North Carolina. The

majority of the preserve (about 85%) supports pocosin

and pond pine woodlands on organic soil (Figure 1).

Only a small portion of the preserve has mineral soils

that support longleaf pine communities or loblolly pine

(Pinus taeda) plantations (currently being restored back

to longleaf pine communites). Fire and hydrology have

shaped the longleaf pine/pocosin (shrub bog)

ecosystems in the coastal Carolinas, including the Green

Swamp, and creating assemblages of species

(communities) with specific components and

characteristics (Christensen 1981). For example, the

Green Swamp is renowned for the high diversity of

herbaceous species in savannas where more than 30

species per square meter can occur (Walker and Peet

1983), including endemics such as Venus flytrap (Dionea

muscipula) and rough-leaf loosestrife (Lysimachia

asperulifolia). Frequent fires (1–10 year return intervals)

are critical to maintaining the open character of the

savannas (Christensen 1981). These open conditions

also are an important environmental factor for the

health and persistence of associated rare species such as

the red-cockaded woodpecker (Picoides borealis) and

Venus flytrap moth (Hemipachnobia subporphyrea) as

noted by Hall and Schweitzer (1993).

Fire naturally has occurred in the pocosins, occasionally

burning into the peaty soils during drought periods

(Hungerford et al 1998). Pocosin fires open up shrub

areas allowing for a mixed-age mosaic of vegetation with

a variety of herbaceous species, including the federally

threatened rough-leaf loosestrife to become established.

The historic fire regime under which pocosins developed

is estimated between 5 to 30 years (Wells 1946; Frost

1995). Fire also stimulates root sprouting and

reproduction and increases nutrient levels in the

nutrient-poor soils of the pocosin (Schafale and

Weakley 1990). The Green Swamp has a range of high

and low pocosin communities, varying with respect to

time since the last fire, the depth of their peat soils, the

saturation and nutrient content of the soil, and the

species they support. Low and high pocosins harbor

such rare species as Cotton grass (Eriophorum

virginicum), laurel (Kalmia cuneata), rough-leaved

loosestrife, arrow arum (Peltandra sagittifolia), beak rush

(Rhynchospora alba) and a sphagnum (Sphagnum

fitzgeraldii) (Schafale and Weakley 1990).

In the absence of fire, all the Green Swamp communities

lose species diversity and build up accumulations of

fuels that can lead to a risk of more intense and more

damaging wildfires. Moreover, with fire suppression,

hardwood shrubs encroach on the formerly isolated

savannas, shading out other species, with a similar loss

of species that favor open conditions in the pocosins.

Fire Management and Research for Biodiversity in the Green Swamp

Margit A. Bucher1 and Maura E. High1

1

The Nature Conservancy, Durham, NC 27707
112 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

The majority of the Green Swamp last burned during a

1932 wildfire; small portions of the pocosins burned in

1955, 1982, and 1990 as wildfires. The fire frequency on

most of the preserve has been reduced through active

fire suppression and lack of prescribed burning on a

landscape level for the past 50 years. The heavy fuel

accumulation (15-25 tons/acre) in the pocosins

increases the risk of severe fires that could burn deeply

into the peat during drought conditions.

The Nature Conservancy has conducted frequent

prescribed burns in the longleaf pine savannas and pine

plantations close to the highway since the early 1980s.

Prescribed burning of savannas in the Green Swamp has

been based on timber management burns during the

dormant season. By burning within 2 days of a rain

event, when the organic soils are saturated, the pocosins

served as a natural fire breaks rather than establishing

destructive plow lines. Such natural fire breaks avoid soil

disturbance, especially in the ecotone between savannas

and pocosin, where many rare and endangered occur.

Most of the isolated, interior savannas and the pocosin,

however, have burned only accidentally within the past

century (Kologiski 1977, McIver 1981).

Developing a Fire Management Plan

for the Green Swamp Preserve

The Nature Conservancy has become increasingly aware

that the limited burning conducted in the past two

decades was not consistent with stated organizational

ecosystem conservation goals. Therefore, TNC developed

the following fire management goals for the Green

Swamp in 1995: (1) to restore and maintain

communities in a spatially and temporally

heterogeneous pattern using prescribed burns as a

Figure 1.—Soil type and preserve boundary of the Green Swamp,

North Carolina
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 113

management tool to mimic fire regimes under which

these communities evolved while protecting and

promoting the rare species associated with them; (2) to

shift from the traditional dormant season to year-round

prescribed burning, but conducting more burns in the

growing season to mimic wildfires as a key natural

process; (3) to plan and time burns to restore and

maintain the mosaic and diversity of habitats in

different successional stages to support the species

indigenous to the Green Swamp; (4) to expand

prescribed burning to the entire preserve, and include

isolated savannas and pocosins, at least on a

experimental basis; (5) to use research and monitoring

as part of the proposed fire management plan to evaluate

the effectiveness of the prescribed burning regime and

redirect management as necessary to accomplish the

management goals (Bucher and Gintoli 2001).

It became apparent to TNC staff that to effectively

manage the longleaf pine/pondpine ecosystems from an

ecological as well as a logistical perspective requires

prescribed burning in the savannas as well as in

pocosins. Fire is needed in pocosin habitat not only to

maintain its inherent diversity but also to reduce the

threat and risk of wildfires originating on the preserve

and threatening adjacent plantations that could lead to

potential impact of suppression activities requiring

heavy equipment. However, conducting safe prescribed

burns in pocosin requires more information on

prescribed burn parameters than is generally available.

The Nature Conservancy learned that the USDA Forest

Service Intermountain Fire Science Lab (IFSL) was

conducting a nationwide study examining prescribed

burning issues in wetlands. In 1995, TNC developed a

joint venture with the IFSL to use the Green Swamp as

one of the research burn sites. The Green Swamp study

goals are (1) to develop guidelines for prescribed

pocosin burns that would not ignite organic soils and

guidelines that could predict when peat fires would

extinguish; (2) to understand the ecological

consequences of suppressing peat fires versus allowing

them to burn; and (3) to provide suppression agencies

with better tools to predict the probability of severe peat

fires on wildfires. In the course of this joint venture, IFSL

staff gathered pre-burn peat moisture and surface

topography data, sampled surface fuels, and monitored

soil consumption and heat transfer during the burn. The

Nature Conservancy staff gathered pre- and post-burn

vegetation data. In cooperation with the North Carolina

Division of Forest Resources (NCDFR), TNC conducted

two experimental prescribed burns in the Green Swamp

pocosin, on 9 September 1999 and 18 September 1999,

to field test the predictive models for peat ignition

developed by the IFSL, (Figure 2) document ignition

patterns and heat transfer in organic soils and monitor

Figure 2.—Aerial view of two experimental pocosin burns in the Green Swamp.

Only the surface fuels burned in the foreground. The burn unit in the back

shows the peat burn unit one year after a burn.
114 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

vegetative response to peat burns and a surface burns to

improve pocosin vegetation models (Bucher 1998).

These carefully planned and implemented burns

demonstrated that pocosins can be burned safely, even

in drought conditions when a peat fires are more likely

to occur. The data from these burns are currently being

analyzed with published outputs planned by fall 2001.

The two experimental burns (each about 10 acres) have

provided data to customize fuel models and have helped

establish parameters for burn plans, but methodologies

to safely conduct prescribed burns on a larger

operational scale of at least a several hundred acres are

not yet realized.

In attempting to return fire to some of the isolated

savannas, TNC staff observed significant differences in

pocosin fuels and fire behavior in different areas of the

preserve. Occasionally burns would move deeper into

pocosin than anticipated, as in a February 1998 fire,

whose path is shown on Figure 3. Rain was approaching

as the fire burned in the pocosin. The fire was allowed to

extinguish itself in the evening, after a flyover

determined it would remain entirely on TNC property.

Afterwards, TNC staff determined that there was a need

for better vegetation and fuels information. In summer

2000, TNC began a vegetation mapping project of the

Green Swamp and its surroundings with Duke

University and the North Carolina Plant Conservation

program. The goals of the mapping project were to

capture the diversity in pocosin from an ecological as

well as a fuels management perspective and develop

classification methods that could easily be exported to

other sites. We employed a remote sensing technique

using four fundamental variables (biomass,

deciduousness, patchiness and soil type) and 65

vegetation plots to classify 7 types of vegetation in the

Green Swamp: longleaf pine savanna; low-, medium-,

and high-density low pocosin; high pocosin, pond pine

woodland; and Atlantic white cedar (Chamaecyparis

thyoides) forest (Kwasny 2001).

Summer 2001 will be used to ground-truth and correct

the vegetation map. Personnel from IFSL will use the

corrected vegetation map and gather additional fuel

characteristics (such as crown-base height, crown bulk

density, and fuel loading) to develop a fuels map and

customized fuel models to enable the use of the Fire

Area Simulator model (FARSITE). The FARSITE model is

a deterministic fire growth simulator that applies fire

behavior calculations to complex environmental

conditions. It allows fuels, weather, and topography to

vary spatially and temporally and produces maps of fire

growth and behavior. It uses current and published fire

behavior models to predict fire behavior (Finney 1998).

After testing the accuracy of the vegetation maps and

customized fuel models developed for the Green Swamp

using recent fires, TNC hopes to better predict fire

behavior in pocosin for the following: (1) to develop

and test prescribed burn parameters using single source

or single line ignition; (2) to simulate the effects of fuel

mitigation along the preserve boundary that may be

effective in reducing the threat of wildfires and increase

the safety of prescribed burns on the preserve to plan

fuel reduction projects; and (3) to predict potential fire

growth and behavior of wildfires and effectiveness of

suppression efforts. Moreover, TNC now has a

Figure 3.—Prescribed burn that burned into the Green Swamp pocosin and

extinguished itself without suppression action.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 115

permanent weather station in the Green Swamp that

provides data for the national fire danger rating system

through a cooperative agreement with NCDFR. Current

and accurate fuel moisture and weather data will be key

components for fire behavior predictions of future burns

and in case of wildfires.

Conclusions

Managing ecosystems using prescribed fire as a

landscape process requires burning across the

boundaries of natural communities and ecosystems to

create a mosaic of habitats and maintain the diversity

inherent to those systems. But any management plan,

especially one that involves a change of practices, must

be based on good observation and research. Setting

goals and carefully monitoring the vegetation response

and effectiveness of the management activities are

essential to developing science-based, ecological land

management. In its Green Swamp preserve, TNC and its

partners are putting these important principles into

practice.

Acknowledgments

The Nature Conservancy has benefited greatly from the

generous sharing of resources by our partners in this

project. We are very grateful to Gary Curcio, Jim Sain,

Shelton Hooks, Robert Houseman, and Miller Caison at

NCDFR, Roger Hungerford, Jim Reardon, Joe Scott, and

Rick Stratton at IFSL, and Jennifer Kwasny and Patrick

Halpin at Duke University as well as all the TNC staff

and volunteers who have helped with field research and

prescribed burning.

References

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Swamp. Rx Fire Notes Newsletter. 7(1).

Bucher, M. A. and L. Gintoli. 2001. Fire management

plan for Green Swamp Preserve, Brunswick County,

North Carolina. Final Report to The Nature

Conservancy, North Carolina Chapter, Durham, NC.

Kwasny, J L. 2001. Mapping vegetation in Green

Swamp preserve for fuel modeling using remote

sensing techniques. MS thesis, Nicholas School of

the Environment, Duke University, Durham, NC.

Finney, Mark A. 1998. FARSITE: Fire area simulator—

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RM-4. 47 p.

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ecosystems. In Fire regimes and ecosystem

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southeastern marshes, peatlands, and swamps.

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Conference. Tallahassee, FL: Tall Timbers Research

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moths, butterflies, and grasshoppers of four Nature

Conservancy Preserves in southeastern North

Carolina. Report to The Nature Conservancy, North

Carolina Chapter, Durham, NC.

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Fire Ecology Conference. Tallahassee, FL: Tall Timbers

Research Station. 19: 78-91.

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Jennings, R. H. 1989. Analysis of natural fire regimes

of the North Carolina Coastal Plain. MS thesis,

Duke University, Durham, NC.

Kologiski, R. L. 1977. The phytosociology of the Green

Swamp, North Carolina. North Carolina. Agriculture

Experiment Station Tech. Bull. No. 250.

McIver, H. 1981. Green Swamp Nature Preserve,

Brunswick County, North Carolina. Report to The

Nature Conservancy, North Carolina Chapter,

Durham, NC.

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of the natural communities of North Carolina,

third approximation. North Carolina Heritage

Program, Division of Parks and Recreation, NC

Department of Environment, Health and Natural

Resources, Raleigh, NC. 325 p.

Walker, J., and R. Peet. 1983. Composition and species

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116 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Abstract.—Although scientific support for fire as a land

management tool has grown, non-industrial private

forest (NIPF) landowners often fail to burn on their

properties. These lands comprise approximately 70

percent of southern forests, making them critical to the

long-term conservation of wildlife and plant species.

Natural resource professionals must overcome key

constraints to use of prescribed fire on NIPF lands if

certain fire-dependent wildlife are to thrive on private

forests. Results from two surveys suggest that fear of an

escaped fire and related liability issues are the greatest

landowner and manager concerns in North Carolina.

Pro-fire media events and public education may be the

best long-term solutions to increase southern NIPF

landowner use of prescribed fire.

Introduction

Anthropogenic fire has a long history in the southern

United States (Hudson 1982; Pyne 1982). Since their

arrival over 10,000 years ago, Native American Indians

burned southern forests and grasslands to drive game,

improve grazing habitat, clear land, and reduce the

chance of wildfire (Hudson 1982; Pyne 1982; Buckner

1989; MacCleery 1993). European immigrants readily

adopted the Indians’ woodsburning practices to improve

range for cattle, reduce fuel loads, kill chiggers and ticks,

increase visibility of snakes and large predators, and

improve access (Stoddard 1962; Pyne 1982). Much of

the Southeast burned every 1-6 years either at the hands

of humans or from natural lightning ignitions (Frost

1998). These high frequency fires helped form the plant

communities now present in the South (Christensen

1981; Buckner 1989; Frost 1998).

Because of its influence on plant communities, fire has

played a central role in shaping the animal communities

of the South as well. The value of fire as a tool to

improve habitat for game species, such as the northern

bobwhite (Colinus virginianus) has long been recognized

(Stoddard 1935). However, fire protection policies,

implemented in the 1920s, facilitated the decline of

southern fire-dependent plant communities, such as the

longleaf pine (Pinus palustris) forest, and the animals

therein (Brockway and Lewis 1997; Brennan 1991;

Engstrom et al. 1996).

Following declines in wildlife populations, much

research was conducted, that highlighted the importance

of fire to non-game species. Pine-dominated stands

burned on short fire rotations (2-3 years) generally have

a more diverse avian community than pine stands

burned using less frequent fire (Wilson et al. 1995;

Burger et al. 1998). Most bird species present in open

pine-grasslands maintained by frequent fire are of equal

or greater management concern than those that occur in

the absence of fire (Sauer et al. 1996; Brennan et.

al.1998; Burger et al. 1998). Declines in herpetofaunal

abundance and diversity can occur following

replacement of fire-adapted vegetation by fire-intolerant

associations (Russell et al. 1999). Most reptile and

amphibian species of conservation concern in the South,

including gopher tortoise (Gopherus polyphemus),

flatwoods salamander (Ambystoma cingulatum), indigo

snake (Drymarchon corais) and pine barrens treefrog

(Hyla andersonii), prefer habitats maintained by frequent

fire (Means and Moler 1979; Brennan et al. 1998;

Russell et al. 1999). Many southeastern small mammal

species thrive in early- to midsuccessional habitats,

many of which historically were created or maintained

by periodic fire.

Although prescribed fire is recognized as a necessary

habitat management tool for many non-game wildlife

species, acreage burned in the South remains relatively

stable and fire continues to be used on only a small

fraction of NIPF lands (Brennan et al. 1998).

Furthermore, NIPF lands that are burned may be done

so only once in the length of a rotation or using fire

frequencies longer than is desirable for maintenance of

quality wildlife habitat (Brennan 1991; Drake 2000).

Prescribed burning on NIPF lands is hampered by

increasingly restrictive federal air quality standards, high

equipment costs, liability risks, multiple ownership

patterns or small tract sizes, financial limitations, and

lack of landowner understanding of fire’s value

(Johnson 1984; Brennan 1991; Brennan et al. 1998;

Izlar 2000). Public attitudes about fire have been greatly

influenced by decades of fire prevention messages (e.g.,

Smokey Bear) emphasizing the destructiveness of

wildfire (Gruell 1991). Although the anti-fire message

excluded prescribed burning or natural fires, most

citizens were unable to distinguish between good or bad

fire (Little 1993). Media coverage has exacerbated the

problem by dwelling on the sensational aspects of

wildfire and doing little to educate the public on the

benefits of prescribed burning (Gruell 1991).

The Role of Non-industrial Private Forest Lands in the Conservation

of Southern Fire-dependent Wildlife

Christopher E. Moorman1, Peter T. Bromley2, Mark A. Megalos3, and David Drake4

1

Department of Forestry, North Carolina State University,

Raleigh, NC 27695

2Department of Zoology, North Carolina State University,

Raleigh, NC 27695

3North Carolina Division of Forest Resources, Raleigh, NC

27699

4

Rutgers Cooperative Extension, New Brunswick, NJ 08901
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 117

Despite the barriers to prescribed burning on NIPF

lands, there are many reasons why management of these

forests must complement efforts to sustain firedependent ecosystems and their associated wildlife

populations on public lands. Non-industrial private

forests comprise a significant percentage (70%) of all

timberlands in the South (Alig et al. 1990). Therefore,

management efforts limited to public lands may not

provide sufficient area to prevent fire-dependent species

from declining or becoming extinct (Brennan et al.

1998). Populations of fire-dependent wildlife on public

lands isolated by adjacent unburned private forests may

experience negative effects (e.g., reduced access to

resources, genetic deterioration, increased susceptibility

to environmental catastrophes) commonly associated

with habitat fragmentation (Harris 1984; Soulé 1987).

Furthermore, private lands adjacent to public forests

may have dangerously high fuel loads and neighboring

landowners may harbor anti-fire sentiments. Either

could eventually limit the ability of public land

managers to burn. In response to relegating too large a

conservation role to government (i.e., public lands),

Aldo Leopold (1949) wrote, “An ethical obligation on

the part of the private owner is the only visible remedy

for these situations.” Now, more than ever, Leopold’s

words ring true.

We use North Carolina as a case study to help clarify

NIPF landowner attitudes pertaining to prescribed fire.

We synthesize several surveys of North Carolina

landowners and certified burners. Then, using literature

accounts and survey results, we identify obstacles to

burning in the South and the possible implications for

southern fire-dependent wildlife. Finally, we discuss

ways to increase both burning on NIPF lands and public

support of prescribed burning as a tool to maintain and

restore critical non-game wildlife habitats in the South.

North Carolina: a Case Study

Surveys of NIPF Landowners

Seventy-nine percent of North Carolina’s forest occurs

on NIPF lands (Alig et al. 1990), making it imperative

that natural resource managers understand NIPF

landowner attitudes related to forest management issues

and what factors impact their land-use decisions. In

North Carolina, 9% of landowners view wildlife (i.e.,

hunting and fishing) as a primary goal for owning and

managing land, whereas an additional 25% manage

their land for timber while protecting the environment

or improving wildlife habitat and recreational

opportunities (Megalos 2000). However, many North

Carolina NIPF landowners neglect management of their

forests because they own small tracts, they do not know

where to start, or they do not rank forestry as a priority

(Megalos 2000). Ninety-one percent of private

landowners in North Carolina own tracts <50 acres, but

these lands comprise only 28% of the total acres of

private lands in North Carolina (Birch 1997).

A survey conducted in the Sandhills region of North

Carolina identified reasons NIPF landowners fail to

burn (Drake 2000). Of 873 landowners owning 330

acres, 81.5% had never burned their forests, and 68.4%

never plan to burn (Table 1). Fire is a relatively popular

management tool in the Sandhills region, meaning that

even fewer landowners are likely to burn in other areas

of North Carolina. Fear of escaped wildfire (41.5%) was

the primary reason most landowners failed to burn, and

29% of those responding did not ever want fire on their

property (Table 2). These reasons may be related to

liability risks associated with prescribed fire and lack of

understanding of the importance of fire in forest and

wildlife management. The cost of burning was not

included as a choice for landowners and may be an

additional inhibitor to burning activities.

Survey of Certified Burners

In an effort to collect additional information on NIPF

landowner attitudes related to prescribed fire, we mailed

a 5-question survey to 292 burners certified through

North Carolina’s training program. Currently, most

certified burners are employed by the North Carolina

Division of Forest Resources, but some work for other

state agencies (e.g., North Carolina Wildlife Resources

Commission, Corps of Engineers) or conservation

organizations (e.g., The Nature Conservancy). Many of

these professionals are in contact with NIPF owners and

were surmised to have excellent insights into the reasons

landowners do or do not burn.

Sixty-four burners responded with partial or complete

surveys. In the opinion of the burners, site preparation

and fuel hazard reduction were the primary motivations

for NIPF owners to burn (Table 3). Conversely, burning

to increase biological diversity was an unlikely

motivation ranked near the bottom of choices (Table 3).

Certified burners ranked liability and smoke

management concerns as the first and third most

important reasons NIPF landowners fail or hesitate to

burn (Table 4). Interestingly, the fear of losing control of

a prescribed fire, which was the primary reason NIPF

landowners in the Sandhills did not burn, was ranked

second out of 11 choices by certified burners (Table 4).

Table 1.—Prescribed fire frequency on non-industrial

private forest lands in the Sandhills of North Carolina

(Drake 2000)

Frequency Sample Size Percent

Never 847 68.4

1-3 Years 847 9.4

4-6 Years 847 5.0

7-9 Years 847 1.8

3 10 Years 847 2.8

Never, But Plan To In Future 847 13.1
118 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Table 2.—Reasons non-industrial private forest landowners failed to

burn their forests in the Sandhills Region, North Carolina (Drake

2000)

Reason for Not Burning Sample Size Percent

Worried About Fire Getting Out Of Control 626 41.5

Don’t Want Fire On Property 627 29.0

Don’t Know Where To Get Assistance 626 22.8

Don’t Like the Looks 626 10.7

Developed Area Nearby 626 3.4

Other 626 24.0

Table 3.—Motivation for non-industrial private forest landowners to burn

their woodlands in North Carolina, 2000 (scale ranges from -2 to 2: -2 =

highly unlikely; -1 = unlikely; 0 = average; 1 = likely; 2 = very likely)

Reasons Sample Size Weighted Average (-2 - 2)

Site Preparation 61 1.21

Fuel Hazard Reduction 62 0.92

Hardwood Competition Control 62 0.63

Manage Game Animal Habitat 59 0.57

Aesthetics 62 -0.25

Increase Biological Diversity 62 -0.63

Pine Straw Production 61 -0.73

Certified burners recommended reducing landowner

and burner liability, increased cost sharing, and more

flexible smoke management guidelines as the

government actions most likely to increase the use of

prescribed fire on NIPF lands (Table 5). Development of

a pro-fire media campaign, which likely would require

less political activity than the top four approaches, was

ranked fifth (Table 5). Thirty of the 44 burners that

responded to a final open-ended question (What would

you recommend in a more specific way to increase the

use of prescribed fire on NIPF lands?) recommended

either a pro-fire media campaign or landowner

education programs to improve the general public’s

understanding and acceptance of prescribed fire.

Discussion

Implications for Wildlife Conservation

With liability and smoke management concerns

identified as the primary barriers to burning, the future

of prescribed burning, which likely will include equal or

greater regulatory and legal restrictions, is uncertain.

Increasingly restrictive air-quality guidelines in the

future may further discourage NIPF landowner use of

fire and may cut short recovery efforts for endangered

species like the red-cockaded woodpecker (Picoides

borealis) (Achtemeier et al. 1998).

Because ecosystems rarely conform to property lines,

their maintenance or restoration generally requires

coordination among multiple entities, including public

land managers and private landowners (Brunson et al.

1996). Although habitat management on private lands

is critical to the sustainability of all wildlife species,

NIPF lands play an especially prominent role in

maintenance of populations of rare and specialized

wildlife species (e.g., fire-dependent species). Fifty

percent of the country’s threatened and endangered

species are found only on private lands and an

additional 20% spend approximately half of their time

on private lands (Hunt 1997). Presently, the diversity of

NIPF landowner backgrounds and objectives results in

an extremely variable forest landscape across the South

(Sheffield and Dickson 1998). Habitat specialists that

are dependent upon fire-maintained habitats are

especially vulnerable to habitat fragmentation (Hunt

1997). Therefore, the current pattern of constant change

in forest condition from one landholding to the next

most likely favors generalist wildlife species.

Brennan et al. (1998) predict continuing decline of

prescribed burning on NIPF lands and refer to the

impending isolation of fire-maintained habitats as an

ongoing land use experiment. Such uncertainty does not

bode well for fire-dependent wildlife. If suitable habitat
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 119

Table 4.—Reasons that best explain why non-industrial

private forest landowners hesitate or fail to burn in North

Carolina, 2000 (ranked from 1 = major reason to 11 = least

important reason)

Reason Sample Average

Liability Concerns 61 3.23

Worried About Losing Control (Wildfire) 60 3.60

Smoke Management Concerns 61 3.74

Neighbor’s Opposition 61 5.16

Cost 61 5.52

Conflict With Local Development 60 5.75

Limited Burning Days (Weather) 60 6.97

Doesn’t Recognize Ecological Value 60 7.05

Doesn’t Know Where To Get Help 61 7.56

Fire Line Considerations 61 7.80

Doesn’t Like The Looks 61 8.84

Table 5.—Government actions most likely to increase the use of

prescribed fire on non-industrial private forest lands in North

Carolina, 2000 (ranked from 1 = major action to 7 = least

important action)

Reasons Sample Size Average

Reduce Landowner Liability 61 3.75

Make Smoke/Fire Regulations More Flexible 61 3.90

Increase Cost Sharing 61 4.11

Reduce Certified Burner Liability 61 4.18

Develop A Pro-fire Media Campaign 61 4.48

Increase Landowner Education On Fire 61 4.62

Increase Direct Professional Assistance 61 4.67

More Proactive State Agencies 61 5.93

is present only as isolated pockets, dispersal by

individual animals is limited and viable populations of

many species may not be maintained over the long term

(Noss 1991). Many fire-dependent wildlife species

already are in decline or are listed as threatened or

endangered (Brennan et al. 1998). As wildlife habitats

are continually lost to population growth and

urbanization, it will be imperative that NIPF landowners

improve management of their forests for wildlife.

Possible Solutions

The problems confronting prescribed burners across the

South are similar to those facing North Carolina’s land

management professionals. Average private landholding

size continues to decline (Birch 1997) and forested

tracts are progressively more isolated from one another

by urban sprawl. Southern NIPF landowners

increasingly are urban and absentee and have multiple

management objectives (Boyce et al. 1986; Izlar 2000).

Frequent changes in parcel ownership inhibit formation

of productive relationships between local professionals

and landowners and make aggressive marketing of state

cost-share and assistance programs imperative. Smoke

management regulations and tort liability add fuel to

the problem by helping to deter NIPF landowners that

otherwise might consider fire as a management tool

(Brennan et al. 1998; Izlar 2000).

Southern resource professionals and landowners agree

that liability concerns and increasingly restrictive air

quality and smoke management guidelines are the

greatest barriers to prescribed burning. Many southern

states, including North Carolina, have passed legislation

that in combination with certified burner training helps

reduce burner liability (Achtemeier et al. 1998). Further
120 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

reduction of burner liability may be unrealistic for

accountability and legal reasons. Rather, legislators,

resource managers, stakeholders and the Environmental

Protection Agency should work jointly on legislation to

exempt prescribed fire smoke emissions from air

quality standards developed to reduce emissions from

cars and industrial smokestacks. Additionally,

continuing education and updates on new technologies

(e.g., fire behavior models, weather prediction models

and risk indices) should lessen the chance of negligent

decisions by burners while widening the prescription

window and increasing the number of acceptable

burning days (Johnson 1984; Lavdas 1996; Achtemeier

et al. 1998).

Divisions within and among agencies often arise

between those who suppress fire and those who use it

(Johnson 1984). To eliminate this dichotomy, the heads

of state agencies must cooperatively lead a more

proactive approach in marketing the benefits of

prescribed fire to those that oppose it and budgeting

sufficient resources to support prescribed burning

efforts. Furthermore, the formation of a prescribed

burning task force in every southern state would allow

transfer of current information among resource

professionals from different agencies and organizations

while promoting constructive discussions on the current

limitations (e.g., air quality guidelines) to use of

prescribed fire on NIPF lands. Task force partners should

include employees of the state agency in charge of fire

suppression, members of the state wildlife agency

including both game and non-game biologists, local

Environmental Protection Agency (EPA) staff,

legislators, and other stakeholders.

Adequate information pertaining to the cost

effectiveness of prescribed burning is not available

(Hesseln 2000). Recent research on the economic

income forgone as a result of declining populations of

fire-dependent species, such as the northern bobwhite

(e.g., Burger et al. 1999), is a step in the right direction.

Most research on the cost effectiveness of prescribed

burning, however, has focused on the economic costs of

burning (e.g., cost per acre) while failing to address

social costs (e.g., costs derived from the inconvenience

of smoke emissions), economic benefits (both market

and nonmarket), and risk (Hesseln 2000). The longterm benefits of prescribed burning, including reduced

risk of catastrophic wildfire, increased forage and habitat

quality for wildlife, and enhanced biodiversity, may

exceed short-term costs like reduced air quality,

decreased aesthetics, risk of escape, and inconvenience

from smoke (Hesseln 2000). However, without research

and documentation of its financial efficacy, large-scale

use of prescribed fire will be difficult to market to NIPF

landowners and the general public. Such analyses would

aid government in defining appropriate funding levels

for support of prescribed burning (Hesseln 2000) and

determining the merit of alternative management styles

like fire suppression.

Natural resources historically have been undervalued

(McNeely 1992). Similarly, there are significantly more

disincentives than incentives for NIPF landowner use of

fire. New financial incentives will be required to offset

costs to landowners not currently using prescribed fire

and to help state agencies, already short on money and

manpower, meet increased demands for burning.

Expanded cost-share programs (e.g., free firelines) can

aid resource-limited landowners wishing to burn. Larger

support budgets for professional assistance would

provide the resources (e.g., on-site equipment, burning

crews available 7 days/week and manpower to oversee

permit approval) necessary to meet anticipated

demands. If budgets are limited, cost-share programs

and professional assistance could be prioritized to fund

only burns that improve wildlife habitat or expand

ecosystem restoration projects. Hazard reduction would

be indirectly achieved on these lands. The creation of

the longleaf pine ecosystem Conservation Priority Area

under the Conservation Reserve Program (CRP) is an

example of cost-share program that indirectly increases

the demand for prescribed fire. Landowners receiving

financial assistance for reestablishment of longleaf pine

forests are more likely to use prescribed fire as a

management tool, and, in the case of CRP landowners,

will be required to use fire to remain eligible for

program payments and benefits.

Forest management plans offer plan writers the

opportunity to interact with landowners and discuss

sustainable resource management alternatives, including

the use of prescribed fire (Megalos 2000). Proponents of

prescribed fire from state land grant universities or

conservation organizations could work with consultants,

state foresters, and other plan writers to encourage the

use of prescribed fire. Government support of

professional assistance programs, such as the Forest

Stewardship Program, may help increase the number of

NIPF landowners with management plans, while

indirectly increasing the number of landowners burning

their forests.

Increasing the use of prescribed fire on small NIPF lands

or on forests within the urban-rural interface will be

difficult. Owners of large forested tracts are most likely

to use governmental cost-share money and public or

private technical assistance (Franklin 1990). These

landowners also are more likely to have a written

management plan for their forests than owners of small

parcels (Birch 1997). NIPF landowners with small

acreages often have non-traditional management

objectives (e.g., non-game management, aesthetic

improvement) and generally own lands nearer to urban

areas where neighbor opposition to fire is common.

Owners of forested lands in the urban-rural interface

may prefer a less-intensive, preservationist approach to

land management.

Although small NIPF landholdings may contribute little

to large-scale restoration of fire-dependent ecosystems,
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 121

informing these landowners of the benefits of prescribed

fire through extension education may improve the

potential for future burning on public and larger private

lands (Cortner et al. 1984; Taylor and Daniel 1984). In

North Carolina, 42% of forest ownerships are between

20-100 acres (Birch 1987), making small tracts a

necessary part of pro-fire programs. All voters and

taxpayers, whether they own forestland or not, can

influence fire-related policy, therefore they also should

be targeted in outreach efforts. Workshops, bulletins,

on-site visits, and land management demonstrations,

traditionally administered by Cooperative Extension

Service agents and specialists, should play a significant

role in improving public understanding of fire-related

issues. Many landowners agree to use of specific land

management practices only after they have seen

successful demonstrations (Brunson et al. 1996).

Cooperative Extension provides a well-established link

for information transfer among land grant universities,

government agencies, and NIPF landowners across the

South. Using modern technology, including the

Worldwide Web and two-way video-teleconferences,

foresters and wildlife biologists can reach a greater

number of landowners more quickly and efficiently

than ever (Bardon et al. 2000).

Public education pertaining to prescribed fire should

focus on America’s youth. Children generally are more

open-minded than adults and are starved for new

information on environmental issues. However, many

live in urban areas and will be exposed to fire only

through their classroom studies or what is seen on

television. Proven environmental education programs

offer a balanced, science-based source of information on

the pros and cons of prescribed fire. Several existing

environmental education curricula, including Project

Wild and Project Learning Tree, contain activities related

to fire ecology. Many conservation organizations and

government agencies have completed or are in the

process of developing similar programs. Natural

resource professionals must aid environmental

educators in distributing and teaching a proactive

prescribed burning educational message in classrooms

across the South.

Ultimately, television and news media must be used to

balance the public’s fear or dislike of fire with its

positive effects. On-site telecasts, performed during a

prescribed fire, could provide a dramatic background to

capture viewer attention and allow burners the

opportunity to discuss fire-related issues. Furthermore,

an aggressive, pro-fire media campaign could help

generate support for increased state agency burning

budgets, exemptions to air quality guidelines, and

legislation to reduce landowner and burner liability.

Like firefighters, prescribed burners should be portrayed

as heroes rather than villains (Murphy and Cole 1998).

The old fire suppression message should be

complemented with a new message relayed by new

characters (e.g., Torchin’ “Tom” Turkey and Burnin’

“Bob”white) that promotes the use of prescribed fire for

fuel hazard reduction, wildlife habitat improvement,

and ecosystem restoration.

With the increasing popularity of herbicides, private

industrial landowners continue to diminish the use of

prescribed fire on their forestlands (Brennan et al.

1998). Herbicides applied for weed control and site

preparation sufficiently fireproof plantations, and

burning counters the value of industry’s frequent

fertilizations unless conducted 33 years after application

(Gerhardt 2000). Similar to NIPF landowners, industry

foresters avoid prescribed fire because of smoke

management and liability concerns (Gerhardt 2000).

More importantly, forest industry’s approach to pine

silviculture often is adopted by local NIPF landowners

and forestry consultants. Herbicides often are portrayed

as valuable wildlife habitat management tools and

adequate substitutes for fire (Brennan et al. 1998).

Rather, herbicides are a potential complement to fire

when habitat management is a primary objective

(Brennan et al. 1998; Brockway and Outcalt 2000). We

believe forest industry should include the use of fire on

NIPF lands as an option in their landowner assistance

programs and maintain parcels of burned forests within

corporate landholdings. Judicious use of fire can be

proof positive of their commitment to plant and wildlife

diversity.

As advances in the use of prescribed fire continue,

resource managers must sell its comprehensive value to

each and every NIPF landowner. Aldo Leopold (1949)

criticized conservationists’ tendency to apply economic

value to all things endangered rather than justify

conservation of species as a “biotic right”. Similarly, we

question threats of potential wildfire as the primary

mode to encourage private landowners to use prescribed

fire. Why not sell prescribed fire based on its ecological

values in addition to its role in fuel hazard reduction?

The more scientific information that can be provided

about the consequences of using or suppressing fire, the

better the general public’s political and social decisions

will be regarding the use of prescribed fire (Van Lear

2000).

Prescribed burning on NIPF lands could significantly aid

restoration and maintenance of fire-dependent plant

communities and the wildlife therein while concurrently

reducing hazardous fuel loads. However, burning must

be conducted frequently and during the right times of

year. As ownership tenure shortens and as tract sizes

lessens and becomes more fragmented, proper use of

prescribed fire as a wildlife management tool will

become increasingly difficult. Overcoming the multitude

of barriers to use of fire will require a joint effort by all

those that will benefit, including the general public. We

feel large-scale, collaborative efforts to educate the pubic

and use of new predictive technologies to reduce risk of

escaped fire are the most effective and efficient means to

increase burning on NIPF lands in the Southeast.
122 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

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124 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Abstract.—Prescribed burning is used for many

silvicultural and wildlife management objectives.

However, the use of prescribed burning can be

constrained due to difficulties in obtaining burning

permits, concerns about liability, potential effects of

scorch on growth and survival of crop trees, its

sometimes ineffective results, limited burning days, and

the costs of applying, controlling, and monitoring

burns. For some landowners, herbicides offer a costeffective alternative to prescribed burning for

manipulating plant communities and wildlife habitat,

especially when the boundaries of application are

closely defined and the focus is on individual habitat

components. Although the ecological effects of fire and

herbicides sometimes differ, when used alone or with

other management practices herbicides offer an

opportunity to meet many wildlife management

objectives. In this paper, we discuss and provide

examples of wildlife management objectives that have

been met by using herbicides, and factors that should be

evaluated when considering use of either prescribed

burning or herbicides.

Introduction

Wildlife habitat is “an area with the combination of

resources (like food, cover, water) and environmental

conditions (temperature, precipitation, presence or

absence of predators and competitors) that promotes

occupancy by individuals of a given species (or

population) and allows those individuals to survive and

reproduce” (Morrison et al. 1992:11). Achieving

management objectives for single wildlife species, or

communities of wildlife species often involves

manipulating in space and time the structure,

composition, and distribution of plant communities

and special habitat features such as snags, down and

dead wood, and mast-producting vegetation.

Fire has long been used for managing plant

communities. Native Americans burned forest land

periodically to improve game habitat, facilitate travel,

reduce insect pests, remove cover for potential enemies,

and enhance native food production (MacCleery 1992,

Day 1953). Early European settlers used fire to improve

habitat for livestock and game species such as whitetailed deer (Odocoileus virginianus), wild turkey

(Meleagris gallopavo), and northern bobwhites (Colinus

virginianus) (Komarek 1981). Currently, foresters and

wildlife managers prescribe fire to reduce fuels, prepare

sites for natural or artificial regeneration, control

competing vegetation in mid-rotation stands, control

certain insects and diseases, enhance development of

forage resources, obtain desired structural characteristics

(e.g., development/promotion of herbaceous and shrub

layers), create specialized habitat components (e.g.,

snags and logs), and restore desired plant species

composition in some ecosystems, e.g., longleaf pine

(Pinus palustris).

As recently as 20 years ago, prescribed burning was used

extensively to manage plant communities on private

lands (Mobley and Balmer 1981), which represents the

majority of lands in the United States (USDA Forest

Service 2000). However, in many states the use of

prescribed burning appears to have been relatively stable

or slightly declining over the past 20 years, although

data related to these trends are limited. In a survey of

southern state forestry agencies by the Georgia Forestry

Commission, 6 responding states indicated that the area

burned over the past 2 decades has remained relatively

stable and 2 states reported a significant decline in area

burned (R. Ferris, Georgia Forestry Commission,

personal communication). Trends in states not

responding are unknown. Data from South Carolina

provides an example of a state where the area burned

annually has been slightly declining over the past 20

years (Figure 1). In areas where use of prescribed

burning is constrained or declining, managers have

begun to search for alternative technologies to achieve

wildlife management objectives.

During the latter half of the 20th century, herbicides

emerged as a tool for manipulating plant communities.

Herbicide products (generally the active ingredient and

one or more surfactants mixed in water) are used

extensively to manipulate the species composition and

structure of vegetation in agriculture, along roads and

utility rights-of-way, in urban settings, and in forest

management (Walstad and Kuch 1987, Brennan et al.

1998). However, data describing trends in herbicide use

in forested ecosystems in the United States are limited.

The recent registration of more selective herbicides

increases the potential to use herbicides for achieving

wildlife management objectives, especially when these

objectives cannot be achieved through prescribed

Herbicides as an Alternative to Prescribed Burning

for Achieving Wildlife Management Objectives

T. Bently Wigley1, Karl V. Miller2, David S. deCalesta3, and Mark W. Thomas4

1

NCASI, Clemson SC 29634

2

Warnell School of Forest Resources, University of Georgia,

Athens, GA 30602

3Northeastern Research Station, USDA Forest Service, Irvine,

PA 16329

4Wildlife Intergration, Birmingham, AL 35244
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 125

burning or in forest systems where fire is not a natural

ecological force. In this paper, we discuss the use of

herbicides to address wildlife-related objectives within

forested ecosystems, with an emphasis on the eastern

United States. We will describe herbicides commonly

used in forest management, silvicultural objectives for

which they are used, habitat components affected, and

wildlife objectives that can be met with their use. We

also will discuss considerations for determining which

tool to use. Our manuscript benefitted from reviews by

R. A. Lautenschlager and D. H. Van Lear.

Ecological Functions of Fire

Can the judicious use of herbicides lead to conditions

similar to those created by fire? The answer depends

upon the specific ecological response in question. Fire

has myriad effects in forested ecosystems. Fire influences

plant and animal species richness, plant reproduction

and development, insect outbreak and disease cycles,

wildlife habitat relationships, soil functions, and

nutrient cycling (SNEP Science Team and Special

Consultants 1996). The ecological effects of fire (Figure

2) are complex, interrelated, and sometimes undesirable

when fire is intense or occurs across large areas (Agee

1993). Pyne et al. (1996), based on information in

Wright and Heinselman (1973), suggested that

depending upon intensity fire may:

• Trigger the release and germination of seeds in some

plant species;

• Stimulate flowering and fruiting of some shrubs and

herbs;

• Alter seedbeds by removing litter and humus and

creating bare soil;

• Stimulate vegetative reproduction of woody and

herbaceous species through overstory reduction;

• Temporarily reduce competition for moisture,

nutrients, and light, thereby favoring some species;

• Selectively eliminate part of a plant community;

• Influence community composition and successional

stage; and

• Regulate susceptibility of forests to blowdowns.

Fire has countless other ecological effects some of which

depend upon the ecosystem in which it occurs. Fire may

kill or injure above- and below-ground portions of

plants, volatize nitrogen, improve conditions for

nitrogen mineralization, cause elements/nutrients to

become more available for uptake by plants, and

dramatically change micro-climates (Wright and

Heinselman 1973). In oak ecosystems, fire creates

favorable conditions for acorn caching by squirrels

(Sciurus spp.) and blue jays (Cyanocitta cristata), reduces

populations of insects that prey on acorns and young

oak seedlings, xerifies mesic sites through consumption

of surface organic matter and exposure of the soil to

greater solar radiation, and reduces understory and

midstory competition from fire-intolerant species (Van

Lear and Watt 1993). Fire scarifies the seed coat of some

plants and enhances their germination, and reduces

debris loading following natural disturbance or

harvesting.

Clearly, use of herbicides also results in some of these

ecological effects. Herbicides can injure or kill the

above-ground portion of plants, selectively eliminate

part of a plant community, influence community

composition and successional stage, and temporarily

reduce competition among plants for resources. In such

cases, herbicides may provide an appropriate substitute

for prescribed burning. However, herbicides cannot

perform every ecological function of fire. For instance,

herbicides cannot directly and immediately alter a

seedbed by removing litter and humus and creating bare

soil, although herbicides can contribute to this

indirectly over time. Herbicides cannot scarify

leguminous seeds to enhance germination or stimulate

seed release in plants such as jack pine (Pinus banksiana).

Herbicides may be more effective at eliciting some

ecological effects if used in combination with other

management tools. For example, mechanical site

preparation could be used in combination with

herbicides to remove litter and humus and create bare

soil. Herbicides and fire already are commonly used in

combination for site preparation to reduce debris

loading and control competing competition. However,

approaches for combining herbicides and other tools to

meet wildlife and ecological objectives need more

thorough investigation.

Herbicide Use in Forestry

Wildlife habitat management is commonly achieved in

conjunction with or as a corollary of other land

management activities such as forestry. Often, the

decision of whether to use fire or herbicides for wildlife

0

50,000

100,000

150,000

200,000

250,000

300,000

1973-741975-761977-781979-801981-821983-841985-861987-881989-901991-921993-941995-961997-98

Year

Hectares Burned

Figure 1.—Area treated with prescribed fire in South

Carolina during 1973-1999 (source, South Carolina

Forestry Commission annual reports).
126 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Figure 2.—The effects of fire on vegetation, soils, hydrology, and geomorphic

processes (from Swanson 1981 and Agee 1993). Excerpted from Fire Ecology of

Pacific Northwest Forests by James K. Agee. Copyright © 1993 by James K. Agee.

Reprinted by permission of Island Press, Washington, DC and Covelo, CA.

management in forested ecosystems depends upon

which tool is most effective at achieving other

landowner objectives (e.g., a forestry objective). Because

herbicides are increasingly a preferred tool for achieving

forestry objectives, we will briefly describe forestryrelated uses of herbicides. Each of these forestry-related

uses represents opportunities for biologists to interact

with foresters and discuss modifications to herbicide

prescriptions that would also achieve wildlife

management objectives.

Herbicides are used in forestry for site preparation,

release of crop trees from competition with herbaceous

and non-commercial woody plants, and timber stand

improvement (Lautenschlager 2000). The reduction of

competing vegetation can significantly increase tree

growth well into mid-rotation (e.g., Zutter and Miller

1998), and controlling both woody and herbaceous

vegetation provides the greatest increase in tree growth

(Figure 3). Herbicide applications typically are tailored

according to soils, structure and composition of the

plant community, and management objectives. Table 1

provides an overview of herbicides commonly used in

forest management.

Depending upon topography and soil conditions, site

preparation may be accomplished using herbicides

alone or in combination with mechanical methods or

fire. When applied for site preparation, herbicides

generally are broadcast. Thus, using herbicides alone for

site preparation (especially when they are aerially

broadcast) generally results in minimal soil disturbance

and erosion potential.

To control herbaceous vegetation, herbicides often are

broadcast or applied in bands or spots during the first

year or two following stand establishment. Some

herbicides, such as sulfometuron can be sprayed over the

top of the seedlings of selected tree species (e.g.,

southern pines) without adversely affecting their

growth. Following stand establishment and through

mid-rotation, herbicides are commonly used to release

crop trees from the influence of competing vegetation.

Sometimes herbicide applications for this purpose

follow thinnings or precede applications of fertilizer.

As an intermediate treatment (timber stand

improvement), hebicides often are applied to individual

woody stems in the midstory and overstory to improve

the composition, structure, condition, and growth of the

stand. Herbicides can be applied during much of the

year to individual woody stems through injection

(herbicide applied to a wound in the tree bole), basal

spraying (herbicide sprayed at the base of the tree close

to the ground), or soil treatment (herbicide applied to

ground), although there may be some seasonal

constraints on these treatments. Treatment of individual

stems is labor intensive, but the ability to do so provides

significant opportunities for selective habitat

enhancement without impacting the entire plant

community.
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 127

The various herbicides registered for forestry use

typically affect different plant species and species

groups. Some herbicides such as glyphosate are broad

spectrum and affect virtually all plant species, although

timing and application rates may alter selectivity of

many herbicides. Other herbicides are more selective

and affect only certain species or plant growth forms

(Table 2). For example, metsulfuron is highly effective

for controlling plants in the genus Rubus. In contrast,

legumes and Rubus spp. generally are tolerant to

imazapyr (Table 3). Fluazifop-P and sethoxdim are

grass-specific chemicals and have little impact on

broadleaf species, while triclopyr has little effect on

grasses and sedges. Sulfometuron methyl (as Oust®) has

been used in northern hardwood forests to control hayscented (Dennstaedtia punctilobula) and New York

(Thelypteris noveborancensis) ferns but was found to have

no effect on woody plants (Horsley 1988a).

Sometimes different configurations of the same

herbicide have different effects, due either to differences

in the nature of the active ingredient (i.e., ester or

amine) or additives (e.g., type of surfactant). For

example, Miller and Mitchell (1990) found that

applications of triclopyr in the form of Garlon® resulted

in 40-80 percent mortality in dogwoods (Cornus spp.),

while applications of triclopyr in the form of

Pathfinder® resulted in >80 percent mortality of the

same species. This selectivity enhances a manager’s

ability to manipulate plant communities. Of course,

because of this differing selectivity, foresters sometimes

mix two or more herbicides in the same tank to enhance

the number of species controlled during an application.

However, some herbicides are not compatible in tankmixes and the number of species controlled by such

mixtures may actually decrease (Ezell 1998).

Using Herbicides to Meet Wildlife

Management Objectives

Although the ecological effects of herbicides and fire

sometimes differ, herbicides can be used to meet many

wildlife management objectives related to plant species

composition and structure, special habitat features (e.g.,

snags, down wood), and the temporal and spatial

distribution of selected habitat components. In fact,

herbicides are more effective than fire for achieving

some wildlife management objectives and can perform

some functions that fire cannot. Numerous studies have

evaluated the potential of using herbicides for specific

wildlife management objectives (Table 4). In reviewing

many of these studies, Lautenschlager et al. (1995)

suggested that, by choosing appropriately (active

ingredient, time of application, application technique),

herbicides can be used to: (1) reduce densities of

invading non-native plants (restoring native populations

and associated wildlife); (2) create snags, dead and

down woody material, and “drumming logs” in early or

later successional stands (providing “old growth”

characteristics); (3) create small, intermediate, or large

early-successional openings within older vegetation

types; (4) change shrub-dominated areas to earlier

successional grassy, or herb/grass-dominated

communities; (5) favor male aspen clones; (6) release

patches or expanses of conifers; and (7) keep woody and

herbaceous “browse” within reach of browsing animals.

Managing Vegetative Species Composition

and Structure

By using newer, more selective herbicides or regulating

time of application, managers can manipulate

understory plant species composition and structure. For

example, dense mats of hay-scented fern and New York

fern can interfere with development of woody seedlings

and the shrub layer in northern hardwood forests

(DeGraaf et al. 1992, Horsley 1988b). This reduces food

resources (fruits from shrubs, woody browse from

seedlings) and vertical structure (shrub and midstory

layers) for many wildlife species, especially songbirds.

Applying herbicides during late summer and early fall

generally will control ferns and result in little if any

damage to desirable woody seedlings or to spring

ephemeral herbs, which already have completed their

annual reproductive cycles and senesced (Ristau and

Horsley 1999)

Figure 3.—Gains in average loblolly pine height, dbh,

stand basal area, and stand volume index over notreatment control through 11 growing seasons by

vegetation control method (from Zutter and Miller

1998). Reprinted from the Southern J. Appl. For.

22[2]:93 published by the Society of American Foresters,

5400 Grosvenor Lane, Bethesda, MD 20814-2198. Not

for further reproduction.
128 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 Table 1.—Characteristics of common silvicultural herbicides and herbicide formulations Herbicide Trade Name Site Conifer Herbaceous Tree Cut Stump Hardwood Basal Bark Activity Behavior Toxicity Preparation Release weed control Injection Application Weed Control Applications in soil Dicamba Vanquish X X X Foliar; Weakly Low Soil adsorbed Fluazifop-P Fusilade DX X Foliar Readily Very low adsorbed, low mobility Glyphosate Accord, X X X X X Foliar Rapidly Very low Roundup adsorbed to soil Hexazinone Pronone 10G, X X X Soil Relatively Low 25G, and MG mobile Hexazinone Velpar L X X X X Soil; Relatively Low Foliar mobile Hexazinone Velpar ULW X X Soil; Relatively Low Foliar mobile Imazapyr Arsenal AC X X X X X Foliar; Weakly Very low Soil adsorbed to soil Imazapyr Chopper X X X X X Foliar; Weakly Very low Soil adsorbed to soil Imazaquin Scepter X X Foliar: Weakly Verylow Soil adsorbed in high pH soils Metsulfuron Escort X Foliar; Moderately Verylow Soil mobile Pendimethalin Pendulum X X Soil Strongly Very low adsorbed to soil Picloram Tordon K X Foliar Weakly Low adsorbed by clays Picloram Pathway X X N/A Weakly Low + 2,4-D adsorbed by clays Picloram Tordon 101M X X X Foliar Weakly Low + 2,4-D adsorbed

by clays

Continued
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 129 Sethoxdim Vantage X Foliar Adsorption Very low varies with organic material Sulfometuron Oust X X Foliar; Not tightly Very low Soil bound (especially high pH) Triclopyr Garlon 3A X X X X Foliar Not tightly Low bound Triclopyr Pathfinder II X X N/A Not tightly Low bound

Herbicide Trade Name Site Conifer Herbaceous Tree Cut Stump Hardwood Basal Bark Activity Behavior ToxicityTriclopyr Garlon 4 X X X X Foliar Not tightly Low

Preparation Release weed control Injection Application Weed Control Applications in soil
130 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

 spp Vitis

 sp. Abies

 spp. Acer

 spp. Alnus

 spp. Amelanchier

 spp. Betula

 sp. Carpinus

 spp. Carya

 sp. Chamaecyparis

 spp. Corylus

 spp. Cornus

 spp. Crataegus

sp. Diospyros 

 spp. Fraxinus

 sp. Juglans

 spp. Juniperus

 spp. Larix

sp. Liquidambar 

sp. Liriodendron 

 spp. Lonicera

 spp. Morus

sp. Oxydendrum 

 spp. Picea

 spp. Pinus

 spp. Populus

 spp. Prunus

 spp. Quercus

 spp. Rhus

 sp. Ribes

 spp. Rosa

 spp. Rubus

 spp. Salix

 sp. Sambucus

 spp. Toxicodendron

 sp. Tsuga

 spp. Ulmus

Table 2.—Susceptibility1 of selected woody plant species to forestry herbicides. Data obtained from product labels and Miller and Mitchell (1990)

 Woody plant species

Herbicide Trade NameDicamba Vanquish; I SSSSS SSTS T S I SSSI I S SSS SSSS

Banvel

Glyphosate Accord; T I S S T SSSI T SI S I TTSI SI SSSSSTI

Roundup

Hexazinone Pronone T I S T S T S T I S T S TTTSI ST SS S

10G, 25G,and MG

Hexazinone Velpar L and T I S I S I S T I I T I TTTSI SI SS SS

Velpar ULW

Imazapyr Arsenal AC S I I T S S SSSS TTSI SS SSS S TS

Imazapyr Chopper RTU S S S S S SSSS S SMetsulfuron Escort S S S S S S S S S S S S S S S S S SPendimethalin Pendulum T T T T T T T T T T T TTTTTT T TTPicloram Tordon K S S I I S I S I S I S I S S S S SPicloram + 2,4-D Pathway SSSSSSS SS S S SSS SPicloram + 2,4-D Tordon 101M S I S S S S I S S S S I SSSS SS SISethoxdim Vantage; Poast T T T T T T T T TTTTT T TSulfometuron Oust R T R R R T R I ITriclopyr Garlon 3A T I S S I S I T S I S S I S I I S SSSS ITriclopyr Garlon 4 I S S S SI TTI I S S I I SI I T SSSS S

Triclopyr Pathfinder II S S S S SS SS SSSS I I SI SI SS S I

1S=Susceptible species (>80 percent control); I=Marginally or variably controlled species (40-80 percent control); T=Tolerant species (<40 percent control); blank = No data available).
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 131

Table 3.—Plant species that are tolerant to imazapyr or that commonly recolonize a site following an

application of imazapyr (from American Cyanamid Company 1999)

Tolerant Recolonize

Scientific name Common name Scientific name Common name

Amorpha fruticosa Indigo bush Amaranthus hybridus Pigweed

Amphicarpa bracteata Hog peanut Ambrosia artemisifolia Common ragweed

Apios americana Ground nut Ambrosia trifida Giant ragweed

Cassia fasciculata Partridge pea Andropogon spp. Broomsedges

Cassia nictitans Small partridge pea Bidens spp. Beggar ticks

Centrosema virginianum Butterfly pea Callicarpa americana American beautyberry

Cercis canadensis Redbud Campsis radicans Trumpet vine

Clitoria mariana Butterfly pea Ceanothus americanus New Jersey tea

Desmodium nudiflorum Beggarweed Chenopodium album Lambsquarters

Desmodium rotundifolium Beggarweed Croton capitatus Wooly croton

Desmodium tortuosum Florida beggarweed Croton glandulosus Dove weed

Ephrosia virginiana Goats rue Cuscuta gronovii Lovevine

Galactia volubilis Erect milk pea Diodia teres Poor-joe

Indigofera caroliniana Wild indigo Epilobium angustifolium Fireweed

Lespedeza bicolor Bicolor lespedeza Euphorbia corollata Flowering spurge

Lespedeza capitata Roundhead lespedeza Geranium carolinianum Wild geranium

Lespedeza hirta Hairy lespedeza Ipomoea purpurea Morningglory

Lespedeza intermedia Wand lespedeza Mollugo verticillata Carpet-weed

Lespedeza japonica Japonica lespedeza Oenothera biennis Evening primrose

Lespedeza procumbens Prostrate lespedeza Oxalis stricta Yellow wood sorrel

Lespedeza striata Common lespedeza Panicum spp. Panic grasses

Lespedeza thunburgii Thunburg lespedeza Parthenocissus quinquefolia Virginia creeper

Psoralea psoralioides Samson snakeroot Passiflora incarnata Maypop

Rhynchosia reniformis Dollar weed Physalis virginiana Ground cherry

Rhynchosia tomentosa Hairy rhynchosia Phytolacca americana Pokeweed

Robinia pseudo-acacia Black locust Polygonum pennsylvanicum Pennsylvania smartweed

Rubus argutus Blackberry Rhus copallina Winged sumac

Rubus trivialis Dewberry Rhus glabra Smooth sumac

Schrankia microphylla Sensitive briar Rhus radicans Poison ivy

Sesbania macrocarpa Sesbania Richardia scabra Florida purslane

Strophostyles helvola Milk pea Rumex hastatulus Sheep-sorrel

Strophostyles umbellata Trailing wild bean Smilax bona-nox Greenbrier

Stylosanthes biflora Pencil flower Trichostema dichotomum Blue curls

Tephrosia spicata Spike tephrosia Viola spp. Violets

Vicia dasycarpa Narrowlearf vetch Vitis rotundifolia Muscadine grape

Vigna suteola Wild pea
132 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Table 4.—Examples of wildlife habitat objectives achieved or resulting through use of herbicides

Objective Location Citation

Reduce live emergent vegetation in wetlands North Dakota Blixt (1993)

South Dakota Solberg and Higgins (1993)

Reduce abundance of parasites in small mammals Oklahoma Boggs et al. (1991)

Increase selected wildlife foods and cover, Pennsylvania Bramble and Byrnes (1983)

and habitat interspersion

Create snags to accelerate development of Oregon Cole (1996)

old-growth characteristics

Manage hardwood midstory in red-cockaded Texas Conner (1989)

woodpeckers (Picoides borealis) cluster areas Georgia Jones (1992)

Reduce habitat suitability for northern pocket Unknown Engeman et al. (1997)

gophers (Thomomys talpoides) to control damage

to lodgepole pine (Pinus contorta) seedlings

Establish food plots Wisconsin Hamilton and Buckholtz (1953)

Control undesirable emergent vegetation and Georgia Wood et al. (1996)

promote waterfowl food plants in impoundments

Manipulate lesser prairie chicken (Tympanuchus Texas Doerr and Guthery (1983)

pallidicinctus) habitat

Improve elk (Cervus elaphus canadensis) and Colorado Kufeld (1977)

mule deer (Odocoileus hemionus) range

Restore herbaceous understory in pine stands Florida Welch (2000)

managed for northern bobwhite (Colinus virginianus)

Create habitat for cavity-nesting songbirds Kentucky McComb and Rumsey (1983)

Provide openings and increase deer (Odocoileus Oklahoma Thompson et al. (1991)

virginianus) forage

Directed application of herbicides also can be used to

suppress some woody species from the shrub and

midstory layers, thus promoting growth and

development of species with more desirable structural

features. For example, shrub-nesting songbirds prefer the

finer and multiple-branching twigs produced by species

such as American beech (Fagus grandiflora) and birches

(Betula spp.) to the more simplified branching of larger

twigs represented by striped maple (Acer pensylvanicum)

(D. S. deCalesta, USDA Forest Service, unpublished

data). Herbicides can be used to selectively reduce the

abundance of striped maple in the shrub and midstory,

which competes with species such as beech and birches.

Such application can be expensive, however, and

uneconomical when treatment levels exceed 400 stems

per acre (R. D. Nyland, State University of New York,

School of Environmental Science and Forestry, personal

communication).

Annual or biennial prescribed burning during the

dormant season has been unable to effectively control

understory hardwood invasion in some open pine

forests managed for red-cockaded woodpeckers (Picoides

borealis) and northern bobwhite (Colinus virginianus).

Welch (2000) reported that a one-time application of

imazapyr alone or combined with prescribed burning

could significantly reduce hardwood invasion without

adversely impacting habitat conditions or food

production for northern bobwhites and allow future

management with prescribed fire during the growing

season. This strategy sometimes is used by federal

agencies (Ralph Costa, U.S. Fish and Wildlife Service,

personal communication).

Herbicides can be used in conjunction with

regeneration techniques, such as a shelterwood harvest,

to alter overstory species composition and structure
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 133

through management of advanced regeneration. A

combined shelterwood harvest and herbicide

application increases sunlight to the forest floor and

stimulates germination and growth of seeds thrown by

the overstory. If conditions are appropriate, prescribed

burning also can be used for similar purposes (Brose et

al. 1999).

Managing Special Habitat Features

The availability of snags and coarse woody debris is a

key factor influencing the abundance and composition

of wildlife communities. Snags and down wood are

created by a number of factors, including shearing

winds, rot associated with insect and disease attack,

lightning strikes, and wild fire. However, these natural

processes produce somewhat variable and unpredictable

results in terms of the abundance and characteristics of

created snags (e.g., species composition, dbh, height,).

Light prescribed burnings may not cause enough

damage to the cambium to lead to tree mortality and

create snags, especially for tree species that are resistant

to fire-induced mortality. Thus, snags and down wood

may not be created from all species.

Herbicides have been used to create snags for a variety of

wildlife-related purposes (Conner et al. 1983, Bull and

Partridge 1986). Because they can be applied selectively

to individual trees, herbicides can be used to regulate

the species composition, dbh, and height of snags and

resulting logs. Snags created with some herbicides (e.g.,

2,4-D) may decay more rapidly than snags created

through other means such as girdling (Conner et al.

1983; Bull and Partridge 1986). However, ongoing

research in Oregon (Michael Newton, Oregon State

University, personal communication) suggests that the

life span of snags created through mechanical means

(e.g., girdling, topping) and herbicides such as MSMA

and triclopyr can be very similar.

Managing Spatial

and Temporal Arrangement of Habitat

Herbicides can be used to manage the spatial and

temporal availability of habitat, a prime determinant of

the diversity and productivity of wildlife communities

(Morrison et al. 1992). For example, herbicides can be

used to create snags and down wood where desired

within the landscape and in a variety of seral stages.

Managers can use herbicides to retain and regulate the

distribution of conifers in riparian ecosystems in order

to provide nesting and foraging habitat for bird species

such as blackburnian warbler (Dendroica fusca),

Swainson’s thrush (Catharus ustulatus), and Acadian

flycatcher (Empidonax virescens). Herbicides can be used

in selected locations to produce patches of earlysuccessional habitats and change overstory species

composition. Overstory species composition can be

changed directly by killing undesired overstory trees or

indirectly and over a long period of time by altering

species composition of advanced regeneration as

previously discussed.

Considerations When Choosing Between

Fire and Herbicides

As vegetation management tools, herbicides and fire

each have a unique set of advantages and disadvantages.

The decision to use fire or herbicides is complex and

involves many variables. We recommend that biologists

and managers consider the following factors when

deciding when and where to apply these tools.

Effectiveness

Obviously, managers should weigh the relative

capabilities of prescribed burning and herbicides to

achieve desired vegetative conditions. For some

conditions, prescribed burning is most appropriate (e.g.,

promotion of fire-adapted understory vegetation).

Sometimes, however, herbicides can be equally or more

effective at eliciting desired vegetative responses. For

example, herbicides are a unique and effective tool for

accelerating the development of late-successional

habitat, specific old-growth components (e.g., large

snags and logs, large live trees of specified species

composition), and associated wildlife species (e.g., Cole

1996). This can be accomplished by turning some live

overstory trees first into snags of desired species, dbh,

and spatial distribution, and later into logs when they

fall. Herbicides are a unique tool for controlling

populations of some non-native species. For example,

Grilz and Romo (1995) found that smooth brome

(Bromus inermis) was most effectively controlled by

spring burning combined with glyphosate applications.

Herbicides are particularly well suited for regulating

plant communities in early successional habitats where

regenerating trees would be damaged by fire.

Historical Disturbance Regime

In selecting whether to use prescribed burning or

herbicides, managers also should consider disturbance

regimes of the ecosystem being managed. Generally,

prescribed burning is most appropriate in fire-associated

or fire-dependent ecosystems such as pine and oak

ecosystems that historically were disturbed on a regular

basis by non-lethal understory fires (Abrams 1992;

Waldrop and Van Lear 1989). However, even in these

forest types fire was not the only form of historical

disturbance. For example, in southern pine forests,

hurricanes, ice storms, and southern pine beetles

(Dendroctonus frontalis) also helped shape forest

structure, species composition, and habitat for species

such as red-cockaded woodpeckers (Picoides borealis)

(Coulson et al. 1995; Hooper and MacAdie 1995;

Conner and Rudolph 1995). These disturbance factors

created important habitat features (e.g., snags, dead

down wood) not readily created through low-intensity

fires with short return intervals.
134 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

In some forest ecosystems, historical fires affected small

areas, were infrequent, or occured primarily as standreplacing fires or mixed and variable fires (Brown 1994,

Runkle 1985). Historical return intervals of fire in some

forest ecosystems in North America are estimated to be

as long as 500-1,000 years, e.g., northern New England

(800 years), upper elevation conifer forests in eastern

Canada (1,000 years), coastal redwood forests in

California (500-600 years) (Oliver and Larson 1990). In

such situations, prescribed burning may not be the most

appropriate tool for achieving habitat objectives and

could cause damage to trees that are not fire-adapted.

For example, in the Northeast, management for species

such as chestnut-sided warblers (Dendroica pensylvanica),

bluebirds (Sialia sialis), and bobolinks (Dolichonyx

oryzivorus) requires development of early-successional

habitat (Braile 2000). However, before timber harvest

can be used to create these habitats, the density of ferns,

grasses, blackberries (Rubus spp.), and undesirable

woody species often must be reduced to allow sufficient

stocking of advanced regeneration of desirable

(ecologically and commercially) tree species. This

objective may best be achieved using herbicides,

particularly in ecosystems where fire is not the dominant

source of disturbance. Where oaks are not fire-adapted,

managers may choose to promote oak regeneration by

top-clipping oak seedlings and treating the remaining

vegetation with a herbicide such as glyphosate. The topclipped oak seedlings, which will not have absorbed the

herbicide, will sprout and grow vigorously in the absence

of competing vegetation (Wright et al. 1985).

Risk to Other Resources

Managers sometimes choose to use herbicides because

fire can damage other resources. For example, extremely

hot fires can alter the physical properties of soils,

accelerate erosion rates, volatize nutrients, and slow

successional recovery (Pyne et al. 1996, Lautenschlager

et al. 1998). Crown scorch can cause mortality and loss

of diameter and height growth in crop trees (Waldrop

and Van Lear 1984). Johansen and Wade (1987)

reported that even slightly scorched trees showed a 15

percent loss of radial growth. Because managed forests

represent a signficant financial investment, many

landowners are hesitant to risk such losses.

Administrative Considerations

Herbicides may be an appropriate tool if administration

of fire is difficult or impossible. For example, fuel loads

may be extremely high, the location may present

difficulties (e.g., near a highway where smoke would

present a hazard to motorists), or labor to administer

the burn may be unavailable. Increasingly, people live in

or near managed forests (Cohen 2000; Egan and Luloff

2000), and because of complaints about smoke and

concerns about potential damage, managers increasingly

are reluctant to burn or are having more difficulties

obtaining burning permits.

Prescribed burnings that escape control are of special

concern to landowners. For example, the May 2000

“Cerro Grande” fire that destroyed a large number of

houses in Los Alamos and White Rock, New Mexico,

began as a prescribed burn on the north rim of the

Grand Canyon. Private landowners often have been the

target of litigation related to unintended consequences

of prescribed burning, and since the passage of the

federal Tort Claims Act, even federal agencies are not

immune from litigation over such matters. In contrast,

drift of herbicides can be minimized by pre-planning

applications using recently developed modeling tools

such as AgDRIFT® (Teske 2000).

Regulations and guidelines at the local, state, and federal

levels also may constrain a manager’s ability to use fire.

Many states have stringent requirements regarding

weather conditions under which prescribed burning can

and cannot be used. For instance, regulations in Texas

prohibit the use of fire under conditions when smoke

will present a hazard on any “pubic road, landing strip,

or navigable water” or when it will affect a “sensitive

receptor” (e.g., a residence, business, farm building, or

greenhouse) (Texas Natural Resource Conservation

Commission 2000). At the federal level, EPA’s interim

air quality policy on wildland and prescribed fire

(Environmental Protection Agency 1998) also

constrains the use of fire in order to regulate emissions

of particulate matter and visibility impairments in the

156 mandatory Class 1 federal areas (“Areas of Great

Scenic Importance”). Regional haze regulations that

eventually will be promulgaed by EPA may further

complicate prescribed burning. When air quality is an

administrative concern, EPA’s Interim Air Quality Policy

on Wildland and Prescribed Burnings (Environmental

Protection Agency 1998) explicitly states that “chemical

treatments may be appropriate tools.”

Economics

Costs obviously are an important consideration when

selecting a habitat managment tool. Generally,

prescribed burning costs less to apply per unit area than

do herbicides. Average costs in the South during 1998

were $40.97/ha for prescribed burning and $178.70/ha

for herbicide applications (Dubois et al. 1999).

However, several other factors also should be considered

when evaluating the cost of fire and herbicides. Multiple

applications of prescribed burning over years or decades

sometimes are required to achieve the same level of

vegetation control that can achieved with one

application of herbicides (Lautenschlager et al. 1998).

Although liability costs and loss of growth do not occur

every time a forest is burned, they could significantly

affect the cost of prescribed burning in some situations

and were not incorporated into estimates by Dubois et

al. (1999). Even without considering these factors, the

cost of applying fire has increased dramatically relative

to the cost of applying herbicides. A cost index
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 135

calculated by Dubois et al. (1991) for prescribed

burning increased at an average annual rate of 10

percent between 1952 and 1988, over twice the rate for

herbicide applications (Dubois et al. 1999).

Operability at Desired Spatial

and Temporal Scales

In deciding whether to use fire or herbicides, managers

also should consider factors related to time and space.

For example, herbicides can be applied to individual

plants, patches of vegetation within stands, and at the

stand scale or larger. In contrast, fire is most easily

applied at the stand or community levels. Herbicides

sometimes immediately produce desired responses in

plant communities (e.g., reduction of non-native

species), while multiple applications of fire over several

years may be required.

Conclusion

The choice of whether to use prescribed burning or

herbicides for achieving wildlife management objectives

depends upon many factors. For achieving some habitat

objectives, herbicides probably are a preferred or partial

alternative to fire. In other cases, fire is the most

appropriate tool. However, prescribed burning

sometimes cannot or will not be used because of

concern about liability, smoke management difficulties,

availability of labor, limited burning days, or other

reasons. In such cases, herbicides may be the only tool

available and must be used if biologists are to even

partially address a wildlife-related objective. Generally,

herbicides are most useful from a wildlife management

prespective for shaping individual habitat components

in well-defined areas. However, no habitat management

tool, whether prescribed burning or herbicides, is best or

even capable of addressing every wildlife management

objective. Thus, we urge managers to retain access to an

assortment of tools, including herbicides, and to use

them in an integrated fashion.

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Abstract.—The role and impact of fire in Southeastern

ecosystems has changed dramatically from pre-European

and early settlement times through present day.

Regionally, pre-settlement fires were caused either by

Native Americans throughout the year or by lightingcaused wildfires during the growing season. Today,

much of the prescribed burning for forest and game

management purposes occurs during the dormant or

winter season in the South, whereas many ecological

restoration or maintenance fires are conducted in the

late dormant season through the growing season

depending on the region and habitat type. Many bat

species in the Southeast presumably have evolved in firedominated ecosystems with roosting strategies that limit

their vulnerability to fire. Moreover, fire in any season

that causes overstory tree mortality and creates snags

suitable as bat roosts probably provide far more benefit

to bats than do the negative impacts from burning.

Dormant season burning may render tree/foliageroosting bats vulnerable to fire in areas of the deep and

mid-South where winter temperatures force prolonged

periods of inactivity and roosts may be limited in highly

fragmented or intensively managed forests. All bats that

tree roost can be impacted by growing season burning if

non-volant young are present. In the winter, cavedwelling myotids seem the least vulnerable to negative

impacts from wildfire and prescribed burning, although

implications from the dramatic increase in late

dormant-early growing season prescribed burning in the

Appalachians and Interior Highlands to bats are

unknown.

Introduction

Little or no research has been conducted on wildfire and

prescribed burning impacts to bats and bat populations.

Consequently, wildlife managers only can approximate

fire effects on these taxa by examining fire impacts to

habitat components important to bats. Although fire

effects on wildlife and the ecological role of fire in

certain Southeastern plant communities such as longleaf

pine (Pinus palustris) forests are fairly well known

(Landers et al. 1989; Landers et al. 1995; Brennan et al.

1998), only recently have wildlife managers started to

understand even the most basic aspects of bat ecology

and habitat use in the Southeast. For most bats, this

understanding is minimal at best (Wilkins 1987;

Watkins 1972; Shump and Shump 1982a; Shump and

Shump 1982b; Kunz and Martin R. A. 1982; Kunz 1982;

Thomson 1982; Webster et al. 1980; Jones 1977; Fenton

1997; Jones and Manning 1989). Bats are the second

most diverse group of mammals in the Southeast and

Mid-Atlantic with 18 species occurring north of

“tropical” midsection of Florida (Whitaker and

Hamilton 1998). To further complicate any assessment

of the fire impact on bats, life histories for these 18

different species differ dramatically and each species

potentially could be affected in profoundly different

ways.

Prescribed burning benefits to bats in the Southeast and

Mid-Atlantic are attributed to forest habitat

modifications that alter or increase amounts and quality

of roosting habitat, modify or improve foraging habitat

and increase arthropod prey abundance. Within heavily

wooded areas, fires that cause overstory mortality and

create canopy gaps may allow bat species such as eastern

red bats (Lasiurus borealis), big brown bats (Eptesicus

fuscus), or other edge-open ecotone specialist bats to

forage more effectively (Edwards et al. 2000). Also,

depending on fire intensity, tree species susceptibility,

and tree canopy position, potential roost snags or

roosting cavities for big brown bats, northern long-eared

bats (Myotis septentrionalis), Indiana bats (Myotis sodalis)

and evening bats (Nycticeius humeralis) can be created.

However, newly created snags may not be immediately

usable by bats. Burning is not an immediate panacea to

a paucity of bat roosting substrate because cavity roosts

or exfoliating bark roosts can take considerable time to

appear and may not be available for one or more

seasons depending on forest community composition

and regional climate.

Burn Season

Most plausible impacts of growing season burns for bats

are poorly documented. Because there are more species

of bats using the forest and because parturition occurs

for most eastern bats at this time, total mortality could

be greater than for dormant season burns. Snags serving

as bat roosts could be consumed by burning and during

intense fires, roosting bats could be killed. Even without

direct mortality, large-scale habitat alteration or habitat

destruction could be detrimental to resident bats during

a time when females presumably are stressed by the

physiological demands of parturition and lactation. The

life histories of many bat species have several attributes

that could be considered adaptive to growing season

fires. During summer months, bats are able to arouse

quickly as the difference between the ambient

Fire and Bats in the Southeast and Mid-Atlantic: More Questions Than Answers?

Timothy C. Carter1, W. Mark Ford2 and Michael A. Menzel3

1

Department of Zoology, Southern Illinois University,

Carbondale, IL 62901

2

Northeastern Research Station, USDA Forest Service,

Parsons, WV 26287

3

Division of Forestry, West Virginia University,

Morgantown, WV 26506
140 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

temperature and active body temperature of bats is less.

Most bats are quick and highly vagile, flying at speeds >

30 kmph (Patterson and Hardin 1969) so that escape

and relocation to unburned areas easily can occur. Most

bat species utilizing trees and snags have multiple roosts

throughout the forest (Sasse and Pekins 1996; Callahan

et al. 1997; Menzel et al. 1998; Foster and Kurta 1999,

Menzel et al. 2001), providing alternate roosts should

the current roost be destroyed by fire. Lastly, most

eastern bat species are able to carry their young for some

time after they are born (Davis 1970). Conceivably, this

would allow females to relocate their young if their

nursery roost became endangered by fire.

Although not necessarily true for the deep South, in the

mid-Atlantic and mid-South, dormant season burns

occur when there are far fewer bats in the forest than in

the growing season. Most dormant season burns tend to

be controlled prescribed fires with a relative low

intensity as currently practiced for pine plantation

management or bobwhite quail (Colinus virginianus)

management. Overstory mortality generally is low and

snag creation often is limited. These lower intensity fires

also may consume fewer existing snags than would more

intense fires at other times in the year. Nonetheless, bats

using forests burned during winter months may not be

fully adapted to this type of rather “unnatural”

disturbance. During winter months, many eastern bats

hibernate or enter prolonged periods of inactivity during

cold snaps. Bats may take up to 30 minutes to arouse

from hibernation (Thomas et al. 1990), making it

difficult for those not hibernating in more protected

cave or mine refugia to reach a body temperature and

activity level sufficient to escape a fire threatening a

roost. Dormant season burning is most prevalent in the

deep South and the Coastal Plain areas of the midAtlantic where winter temperatures generally are warm

enough to ensure year-round presence of bats. In upland

hardwood forests in the mid-Atlantic states, such as West

Virginia and Virginia, burning for ecological restoration

efforts to promote oak (Quercus spp.) regeneration

(Brose et al. 1999) takes place from mid-April to midMay. During this time, many bat species are finished

hibernating and already could be using these upland

forests for roosting and foraging.

Solitary Foliage-roosting Guild

Bat species in this guild use the forest exclusively for

roosting. Regionlly, this includes the eastern red bat,

Seminole bats (Lasiurus seminolus), and hoary bat

(Lasiurus cinereus) that roost in tree foliage throughout

the year. Eastern pipistrelles (Pipistrellus subflavus) roost

within tree foliage during the summer (Shump and

Shump 1982; Menzel et al. 1998, Veilleux 1999). The

northern yellow bat (Lasiurus intermedius) generally

roosts in Spanish moss (Tillandsia usneoides; Webster et

al. 1980; Menzel et al. 1999). With the exception of

females nursing young, all individuals in this guild roost

solitarily. We hypothesize that these roosting habits

translate into few or minimal fire effects on this group

as disturbance events probably affect fewer animals at

any single time as opposed to colonial-roosting bats.

Furthermore, their habit of foliage roosting permits

quick perception of surroundings and allows for faster

response times. In largely forested landscapes, there are

almost infinite amounts of available roosts for alternate

use. However, in highly fragmented landscapes, fire and

smoke may drive these bats to alternate forest patches as

witnessed by Rodrigue et al. (2001), increasing their

vulnerability to daytime predation as bats cross nonforested areas.

During winter months, this roosting guild is composed

exclusively of Lasiurines, as the eastern pipistrelle

retreats to cave or mine hibernacula. Of bats that

continue forest roosting throughout winter such as the

eastern red bat, some individuals migrate to the deep

South and continue similar roosting and activity habits,

whereas others individuals staying in the mid-Atlantic

and upper South will periodically retreat to the leaf litter

on the forest floor to escape cold snaps. Eastern red bats

have been observed flying up from the leaf litter in front

of dormant season fires (Saugey et al. 1998; Moorman

et al. 1998). Whether or not eastern red bats or any

other foliage roosting bats experience significant annual

mortality from dormant season burning is unknown. An

extensive study of the winter roosting habits and the

possible effects of fire on this roosting guild has not

been conducted.

Bark and Cavity Roosting Guild

This guild uses tree cavities and exfoliating bark for

roosts. Generally, these bat species form small roost

colonies that render them more susceptible to fire effects

than solitary foliage roosting bats. As a result, even a

single, localized fire event potentially could impact more

individual bats. Cavity roosting bats include the big

brown bat, evening bat, Rafinesque’s big-eared bat

(Corynorhinus rafinesquii), northern long-eared bat, and

silver-haired bat (Lasionycteris noctivagans). Regionally,

evening bats often are found roosting in upland pine

forests (Menzel et al., 2001) where both natural and

anthropogenic fires are common. Because these bat

species roost internally within cavities, their perception

and response time necessary for exit and escape may be

delayed. Bark roosting species include the endangered

Indiana bat, little brown bat (Myotis lucifugus), and at

times, the northern long-eared bat (Menzel et al. in

press). In the northern parts of its distribution, the

southeastern bat (Myotis austroriparius) will use

exfoliating bark and hollow trees, whereas in the

southern portion it also utilizes caves, bridges and other

structures as roosts (Gardner et al. 1992). The small to

medium sized maternity colonies formed under

exfoliating bark in trees and snags by this roosting guild

are vulnerable to fire disturbance in several ways. Snag

roosts, depending on age and condition and fire

condition can be highly combustible. However, most of
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 141

these species use multiple summer roosts that combined

with their ability to carry young in flight might offset

this significant liability (Davis 1970). Also, many of

these bats often roost in riparian areas where prescribed

burning generally is not applicable and where wildfire

risks are slight (Gardner et al. 1991; Gardner et al. 1992;

Foster and Kurta 1999).

During winter, all the myotids as well as big brown bats

and eastern pipistrelles in the foliage roosting guild

move to hibernacula in caves, mines, and large

rockhouses and clifflines with cave-like characteristics.

Silver-haired bats winter from the lower Ohio Valley and

mid-Atlantic south into the mid-South where they

switch to rock crevice and solitary bark roosting habits.

Evening bats migrate to the deep South and coastal

regions and continue using tree cavities for roosts. We

hypothesize that fire effects on evening bats and silverhaired bats during winter months would be similar to

those of the summer months with the added caveat that

these animals could be in a prolonged torpor with

slower reaction times.

Cavernous Roosting Guild

Of the bats that use caves and mines, few use

underground roosts during the summer months. Most

use forests (cavities and exfoliating bark) during the

summer months. Bats that summer-roost in caves or

mines include southeastern bats, gray bats (Myotis

grisescens) and Townsend’s big-eared bats (Corynorhinus

townsendii). Bachelor colonies of northern long-eared

bats and Indiana bats occasionally have been found in

caves and mines during the growing season. Although

subterranean roosts offer direct protection from flames,

fire near occupied underground roosts can impact bats

both directly and indirectly. Fire that alters surrounding

vegetation such that cave or mine airflow is modified

potentially could affect entrance and chamber

microclimate (Richter et al. 1993). Additionally, habitat

changes around caves or mines may impact forgaging

bats during pre-hibernation swarming or posthibernation pre-dispersal, two feeding periods when

bats must gain critical fat reserves. Smoke impacts to

cavernous roosting bats depend on a cave’s airflow

characteristics. Depending on the season and air

temperature, cave and mine entrances serve as entrances

and exits for warm air or cold air (Tuttle and Stevenson

1977). A fire that creates smoke up wind from a

“breathing” entrance potentially could fill the cave with

smoke. In the late spring and summer, maternity

colonies of gray bats or Townsend’s big-eared bats that

seek warm areas in a cave could be directly influenced by

smoke drawn into the portions of a cave serving as a

warm air trap or reservoir. During the winter months,

the cavernous roosting guild also expands to include the

rest of the myotids, eastern pipistrelles, big brown bats

and Rafinesque’s big-eared bats. Most caves and mines

used as hibernacula tend to trap colder air in the winter,

so smoke entry into the system probably would be less

likely to occur. Nonetheless, if smoke from a nearby fire

is drawn into the hibernaculum, the potential for a

catastrophic event affecting large groups of bats is a

possibility. Mortality from smoke inhalation or

decreased fitness from premature overwinter

disturbance and arousal are real possibilities. The

increased interest in the use of fire to manage upland

hardwood forests in the mid-South and mid-Atlantic

where karst topography and cave formation occurs

merits expanded fire-ecology research.

Crevice Roosting Guild

Regionally, the myotids, Rafinesque’s big-eared bat, and

Townsend’s big-eared bat occasionally will utilize rock

outcrop and clifflines during the growing season for

roosting habitat. Also, silver-haired bats have been

observed using rock outcrops as winter roosts in

Arkansas (D. Saugey, USDA Forest Service, pers. comm.).

However, during the summer months, the small-footed

bat (Myotis leibii) almost exclusively uses cracks and

chimneys in large rock outcrops and clifflines (Whitaker

and Hamilton 1998). Depending on localized geology,

these roosts can be found in a variety of forested

habitats and conditions in the region. Within the

mountainous portions of the Appalachians in the midSouth and mid-Atlantic, rock outcrops occurring along

xeric ridgelines often are covered by dry-site oak

communities or fire-adapted pitch pine (Pinus rigida)-

table mountain pine (Pinus pungens) overstories with

extremely flammable ericaceous shrub layers. Although

regional interest in prescribed burning in these systems

to maintain these relictual pine types is increasing (Vose

et al. 1997; Welch and Waldrop 2001), the impacts to

bats such as the small-footed bat are unknown.

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144 The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288

Wildlife, Habitat, and Prescribed Fire in the Southeastern

Coastal Plain: Past, Present, and Future

Leonard A. Brennan1

Abstract.—Fire has been an imperative ecological force that has shaped the natural history and

culture of the Southeastern Coastal Plain. In the past, humans have influenced the “ natural” role

of fire in the southeast, first by Native Americans, then by Spanish and other European settlement,

and finally by modern culture. As human culture developed, fire, and how it influenced wildlife

populations on both the landscape and local scales, became more and more removed from the set

of environmental conditions that originally allowed it to shaped the biota of the southeastern

landscape. Presently, broad scale habitat fragmentation from urbanization, agriculture, and

intensive forestry, has eliminated fire from much of the southeast. Also, at the present time,

virtually all of the vertebrates in the southeast that are declining, threatened, or endangered require

open forest conditions maintained by frequent fire. In the future, most fire in the southeast will

probably be relegated to isolated fragments of public lands, ecological preserves, and private

hunting plantations. However, there are some developments that have emerged from research, as

well as management experience and common sense, that have promise. For example, fire ecologists

and managers have learned to pay respect to the use of fire is emerging. Such an approach

transcends traditional single species, game management objectives, and incorporated consideration

of how keystone native plants, arthropods, and other nongame vertebrates, respond to variation in

seasonal fire applications, as well as to variation in fire return intervals.

1

Tall Timbers Research Station, Tallahassee, FL 32312

The Role of Fire in Riparian Zones of the Northern Rocky Mountains

Elaine K. Sutherland1 and Kevin McKelvey1

Abstract.—While the importance of riparian systems in the northern Rocky Mountains as sources of

productivity and diversity is recognized, there is little information about the interaction between

pattern and process. To sustain these areas, we need to understand the characteristics of disturbance

processes and how they result in patterns in these systems. There is mounting evidence that simply

protecting riparian areas from fire and other disturbances results in unsustainable riparian

conditions and deterioration of habitat for wildlife. Recent high-intensity fires, which in the past

were probably uncharacteristic of the upland vegetation surrounding riparian zones, can result in

soil erosion and sediment loading that can damage aquatic systems. However, low-intensity fires

play an important role in determining composition and structure in upland and streamside

vegetation, and consequently in contribution of vegetation debris to the aquatic system. Without

low-intensity fire, uplands and streamsides succeed to shade-tolerant coniferous species, with

consequently reduced dominance or loss of early-successional deciduous trees and shrubs. These

altered conditions can have important consequences for habitats of terrestrial and aquatic fauna.

1

Rocky Mountain Research Station, USDA Forest Service, Missoula, MT 59807
The role of fire in nongame wildlife management and community restoration: traditional uses and new directions GTR-NE-288 145

The Restoration of Oak-Hickory Forests in the Central Hardwoods:

Results of a Landscape-scale Prescribed Burning Experiment in Ohio

Elaine K. Sutherland1 and Todd F. Hutchinson2

Abstract.—Oaks have been an important component of eastern deciduous forest communities since

the end of the last glaciation, 10,000 years BP. There is wide recognition that oaks are highly fire

adapted, and that fire played an important role in the ecology of oak forests in the past, particularly

in promoting the dominance of oak in regeneration layers. In southern Ohio, forest composition

and structure has changed since aggressive fire protection was implemented. Although oaks still

dominate the overstory, advanced regeneration is dominated by species such as yellow-poplar,

maples and blackgum. The objectives of this project are to determine ecological response of mixedoak communities in southern Ohio to prescribed underburning under frequent and infrequent fire

regimes. Four study areas, ~ 90 ha each, were selected in 1994 in the hilly country of the

unglaciated Allegheny Plateau. These areas are dominated by oak forests > 80 years old. Study

areas were stratified by an estimator of available soil moisture based on landscape physiographic

characteristics and soil water holding capacity. This integrated moisture index (IMI) is a good

predictor of vegetation composition, as well as songbird species occurrence. Study areas and plots

were installed in 1994 and pre-burning data were gathered in 1995. The four study areas are

divided in thirds; two early spring burning treatments and an unburned reference. The frequent

treatments were burned annually (1996-1999) and infrequent treatments were burned twice (1996

and 1999). Physical and biological attributes of the forests were fully monitored between 1995 and

1999. These low-intensity fires had significant effects on soil and litter distribution and

composition and structure of herbaceous and small diameter vegetation, but many of these effects

likely will be short-lived. There were consequent changes in ground-nesting bird populations, but

not in those nesting elsewhere. Insect communities were little altered.

1Rocky Mountain Research Station, USDA Forest Service, Missoula, MT 59807

2Northeastern Research Station, USDA Forest Service, Delaware, OH 43015

Fire Ecology of Marshes and Canebrakes in the Southeastern

United States

Cecil Frost1

Abstract.—Most marshes in the United States can be considered fire communities. With the

exception of true salt marsh (salinity 3-4%), maintenance of species diversity in marshes is

dependent upon frequent fire. Plant species diversity in marshes decreases as salinity increases and

increases as fire frequency increases. Canebrake, another fire community, once covered at least 10

million acres in stream bottomlands and peatlands of the southeastern United States. This

comprised critical habitat for a number of rare plant and animal species. Far less than 1% of this

habitat remains, canebrakes having disappeared after fire was removed from the landscape. When

fire is kept out, canebrake succeeds to multistoried wooded communities such as bottomland

hardwoods, pocosin, pond pine forest, red maple forest, and bay forest. Rare, fire-dependent plants

native to canebrakes include golden sedge (Carex lutea), Lilium iridollae and Lilium gazarubrum.

Important wildlife species include Swainson’s warbler, Bachman’s warbler, and a butterfly, St.

Francis satyr. Canebrake also is a fire frequency indicator community. A historical record of

canebrake for a particular location indicates that that region once experienced a frequent fire

regime. Studies of succession in canebrakes indicate that the original fire frequency ranged mostly

between 2-8 years between fires. Protection and restoration of canebrake requires management

with fire, and has been almost entirely overlooked in site management and conservation planning.

1

North Carolina Plant Conservation Program, Department of Agricultural and Consumer Services,

Raleigh, NC 27611
Printed on Recycled Paper

Ford, W. Mark; Russell, Kevin R.; Moorman, Christopher E., eds. 2002.

Proceedings: the role of fire for nongame wildlife management and

community restoration: traditional uses and new directions; 2000

September 15; Nashville, TN. Gen. Tech. Rep. NE-288. Newtown Square, PA:

U.S. Department of Agriculture, Forest Service, Northeastern Research Station.

145 p.

Contains 16 papers and 4 abstracts on the use of fire for nongame wildlife

management and rare plant community management in restoration in the United

States with an emphasis on forested systems.

Keywords: alternative management, community restoration, history of fire,

prescribed fire, nongame wildlife
Headquarters of the Northeastern Research Station is in Newtown Square,

Pennsylvania. Field laboratories are maintained at:

Amherst, Massachusetts, in cooperation with the University of Massachusetts

Burlington, Vermont, in cooperation with the University of Vermont

Delaware, Ohio

Durham, New Hampshire, in cooperation with the University of New Hampshire

Hamden, Connecticut, in cooperation with Yale University

Morgantown, West Virginia, in cooperation with West Virginia University

Parsons, West Virginia

Princeton, West Virginia

Syracuse, New York, in cooperation with the State University of New York,

College of Environmental Sciences and Forestry at Syracuse University

Warren, Pennsylvania

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