United States

Department of

Agriculture

Forest Service

Northeastern

Research Station

General Technical

Report NE-274

Proceedings: Workshop on

Fire, People, and the Central

Hardwoods Landscape

March 12-14, 2000

Richmond, Kentucky
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Proceedings: Workshop on Fire,

People, and the Central

Hardwoods Landscape

March 12-14, 2000

Richmond, Kentucky

Compiled by:

Daniel A. Yaussy

Sponsors

Eastern Kentucky University, Division of Natural Areas

University of Kentucky, Department of Forestry

University of Tennessee, Department of Forestry, Wildlife & Fisheries

USDA Forest Service,

Daniel Boone National Forest

Northeastern Research Station

Region 8

Region 9

USDI Fish and Wildlife Service

National Park Service, Great Smoky Mountains

Kentucky Division of Forestry

Kentucky Department of Fish and Wildlife Resources

Kentucky State Nature Preserves Commission

Forest Landowners Association

Kentucky Forest Industry Association

The Nature Conservancy

Kentucky Woodland Owners Association
Preface

This conference was conceived as a vehicle to demonstrate to forest land managers and the

interested public, the historical role fire has played as an agent of change and renewal in forests

of the area known as the Central Hardwoods Region. However, fire is widely perceived as being

potentially dangerous and damaging to trees and ecosystems. This conference, held in

Richmond, Kentucky, March 12-14, 2000, explored the historical, present, and future roles of fire

and humans in shaping the vast and diverse deciduous forest of the Central Hardwoods Region.

This is a compilation of the 18 papers and 16 poster abstracts presented in the one and a half

days of the conference. The workshop was followed by a field trip to Cliff Palace Pond where the

pollen and charcoal record evidence of the relationships of fire, people and the forest that

existed more than 9500 years ago, challenging our view of the “virgin” forest primeval. A few

speakers chose not to submit a final paper to this compilation and the abstract of their

presentation is included here.
Table Of Contents

Presentations

Introduction

Fire in the Central Hardwoods: Why are We Concerned? ................................................................................ 1

Rex Mann

Keynote Address – Humans and Fire, A History of the Central Hardwoods ..................................................... 3

Ray Hicks

The History of Fire

Native Burning in Western North America: Implications for Hardwood Forest Management.......................... 19

Charles Kay

Humans, Topography, and Wildland Fire: The Ingredients for Long-term Patterns in Ecosystems ................ 28

Richard Guyette and Daniel Day

Fire Ecology

Fire on the Edge: Prehistoric Fire along the Escarpment Zone of the Cumberland Plateau .......................... 36

Cecil Ison

Fire and the Ecological History of Oak Forests in the Eastern United States ................................................. 46

Marc Abrams

Effects of Fire on the Ecology of the Forest Floor and Soil of Central Hardwood Forests.............................. 56

Ralph Boerner

Fire and Understory Vegetation: a Large-scale Study in Ohio and a Search for General Response

Patterns in Central Hardwood Forests ............................................................................................................ 64

Todd Hutchinson and Steve Sutherland

Fire and Birds in the Central Hardwood Landscape ....................................................................................... 75

Todd Engstrom

Stream Mayflies, Stoneflies, and Caddisflies: Do They Care About Controlled Burning in the Forest? ......... 87

Jack Feminella

Fire Bugs ........................................................................................................................................................ 88

Jim Bess

Black Bear Ecology and the Use of Prescribed Fire to Enhance Bear Habitat ............................................... 89

Keith Weaver

Fire and Ecosystem Management

Using Prescribed Fire to Regenerate Oaks .................................................................................................... 97

David Van Lear, Patrick Brose, and P.D. Keyser

Current Research on Restoring Ridgetop Pine Communities with Stand Replacement Fire ....................... 103

Thomas Waldrop, Nicole Turrill Welch, Patrick Brose, Katherine Elliott, Helen Mohr,

Ellen Gray, Frank Tainter, Lisa Ellis

Barrens Management and Restoration: A Kentucky Example .......................................................................110

Joyce Bender
Fire and the Future Forest

Resistance is not futile: The response of hardwoods to fire-caused wounding ............................................. 111

Elaine Kennedy Sutherland and Kevin Smith

Managing Rare Plant Populations with Fire in Great Smoky Mountains National Park .................................116

Janet Rock

In Summary: Fire in the Evolution of the Eastern Landscape: A Timeline .................................................... 120

Ed Buckner

Poster Abstracts

Oak Seedling Response to Fire and Herbivory ............................................................................................ 122

A.S. Adams and L.K. Rieske

Prescribed Burning: Effects on Forest Bird Populations ............................................................................... 122

Vanessa L. Artman

Ecological Effects of Fire on Xerophyllum Asphodeloides, A Rare Appalachian Lily .................................... 123

Norman A. Bourg and Douglas E. Gill

A Shelterwood–burn Technique for Regenerating Productive Upland Oak Sites ......................................... 123

Patrick Brose

Effects of Fire, Deer Herbivory, Seed Predation, and Canopy Gaps on Forest Regeneration:

A Large-scale Experimental Approach .......................................................................................................... 124

Rachel J. Collins and Walter P. Carson

Effects of Shelterwood Harvesting and Prescribed Fire on Regeneration Structure .................................... 124

Wade Dorsey and Mary Ann Fajvan

Prescribed Fire Effects on Foliar Nutrients, Photosynthesis, and Growth of Ridgetop Maple and

Oak Seedlings .............................................................................................................................................. 125

N.L. Gilbert, M.A. Arthur, S.L. Johnson, S.K. Gleeson, and B.A. Blankenship

Chilton Creek: A Case Study of Landscape Fire Effects ............................................................................... 125

George W. Hartman

Ethnoecology of Fire: An Experimental Approach in the Ohio Valley............................................................ 126

Erin B. Hines, Jan Salick, Elaine Kennedy Sutherland, Todd F. Hutchinson, and Steve Sutherland

Managing Red-Cockaded Woodpeckers (Picoides Borealis) Affects Breeding Bird Communities of

Pine-Oak Forests in Southeastern Kentucky ................................................................................................ 126

Martina Hines

Disturbance History of an Old-Growth Forest in Southeastern Ohio ............................................................ 127

Brian C. McCarthy and Darrin L. Rubino

Response of Pin Cherry (Prunus Pensylvanica L.F.) to Fire, Canopy Disturbance, and Deer Herbivory

on the Westvaco Wildlife and Ecosystem Research Forest ......................................................................... 127

David W. McGill, Rachel J. Collins, and Walter P. Carson

Prescribed Fire and Herbicides Modify Soil Processes During Barrens Ecosystem Restoration ................. 128

C.C. Rhoades, T. G. Barnes, and B.E. Washburn

Experimental Studies of Oak Seed Responses to Fire Temperature in Appalachian Mixed-oak Litter......... 128

Cynthia L. Riccardi and Brian C. McCarthy

Monitoring Scarab Populations Exposed to Prescribed Fire in Mixed-oak Forests of Southeast Ohio ........ 129

A.E. Smith and D.J. Horn

Ground Beetle Responses to Prescribed Burning in Southern Ohio’s Hardwood Forests ........................... 129

Chris Stanton, Foster F. Purrington, and David J. Horn
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 1

Abstract

For several decades, land managers in the Central

Hardwood Region have watched their oak-dominated forests

change in the absence of fire. Stand structure has changed

as a dense understory of shade-tolerant, fire-intolerant

species developed. The forests were not sustaining

themselves.

This same phenomenon was occurring in many parts of the

United States, resulting in forest health concerns. Severe

wildfires erupted in places like Yellowstone National Park

and Florida.

Evidence is accumulating that the oak, chestnut, and yellow

pine plant communities in the Central Hardwood Region are

the result of centuries of burning by Native Americans. To

maintain the diversity of our forests, fire must be restored to

its vital role.

There are a number of myths relating to the forests that are

widely held by the public that present obstacles to

reintroducing fire. These myths must be dispelled if we are to

restore fire to its rightful role and thus sustain our forests.

Introduction

For years, land managers throughout the Central Hardwood

Region have noted definite changes occurring throughout

the historically oak-dominated forests under their care. The

oaks, which are valued because of their usefulness as

wildlife food sources and for a variety of wood products, are

being replaced with maple, hemlock, white pine, and other

species. Oak seedlings are simply not surviving in the dense

understory shade beneath the oak stands. In the absence of

fire, the maple, hemlock, white pine, and other species have

come to dominate the understories and are posed to capture

the overstory.

Indeed, the only way we could regenerate the oak forest was

to harvest it and manage the coppice growth. About the only

place acorns were germinating and growing to mighty oak

trees was at the edge of the forest, where sunlight was

abundant. In addition, an ever-increasing number of plants

and animals were being listed as endangered or threatened.

Something was missing here. Yet, in view of the amazing

recovery our Eastern forests made from the massive

timbering that occurred during the early 1900’s, we were not

terribly concerned until fairly recently.

Throughout many other parts of the country, similar

challenges have emerged—challenges to restore firedependent forest ecosystems that are being replaced in the

absence of fire. These conditions are a result of more than

75 years of fire suppression and in some areas have been

labeled as forest health problems.

For those of us who serve as firefighters, we have had

numerous opportunities to witness the day of reckoning that

eventually comes to fire-dependent ecosystems when

historic fire is suppressed for long periods of time.

Yellowstone in 1988, and more recently the Florida fires of

1998, were classic examples of forest succession in the

absence of “normal” fire. Fuel loads and more volatile

undergrowth in some areas of the country have increased in

the absence of fire to a point where even carefully applied,

controlled burns cannot be ignited without disastrous results.

Indeed, there will be plenty of gainful employment for

wildland firefighters as these past-due bills continue to come

due in the years and decades ahead.

Closer to home, in the Central Hardwood Region, evidence

is accumulating that our forests are also fire-dependent

communities. Consensus is emerging that Native American’s

fire was responsible for the dominance of the oaks, American

chestnut, and yellow pine plant communities we have

enjoyed in recent history. Even though wildfire hazards in

this region are not nearly as severe as those in the coastal

plains or in the West, the exclusion of fire has greatly altered

the future of oak and pine-dominated ecosystems. The oak

and pine forests are not sustaining themselves in the

absence of frequent surface fires except where land

managers have interjected silvicultural manipulation.

As we continue to learn more about the ecosystems that

make up the Central Hardwood Region, and as we continue

to sort out the relationships between individual species of

plants and animals and their ecosystems, there appears to

be a common thread linking many of these problems we

face—the exclusion of frequent surface fires.

As we learn more about these forests and how they came to

be, we are dispelling more and more the myths that we as a

people have come to believe. For example,

• There is the myth of the pristine wilderness that existed at

the time of European settlement, where a squirrel could

theoretically travel through the unbroken treetops of an oldgrowth climax forest from the Atlantic coast to the Mississippi

River.

• There is the myth that Native Americans were few in

number and had little impact on the land.

• And there is the myth that the forest was very stable, that

disturbances were small in scope and natural in origin, and

that a great balance existed.

Fire In The Central Hardwoods: Why Are We Concerned?

Rex B. Mann1

1

Timber/Wildlife/Fire Staff Officer, USDA Forest Service, Daniel

Boone National Forest, 1700 Bypass Road, Winchester, KY

40391
2 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Now we know that the Central hardwood landscape at the

time of European settlement was very diverse with many

treeless openings. The pristine wilderness and unbroken

canopies existed only in the myth. It is also generally agreed

that Native Americans were far greater in numbers than we

have been led to believe, until European diseases decimated

them. We also know that the tremendous landscape diversity

we inherited was a result of their land use, primarily through

their use of fire. And finally, we now know that the great

“balance of nature” did not exist, that change was the norm

rather than stability.

As the land-managing agencies and land owners enter this

new millennium, we are challenged to look into and

understand the past in order to be able to craft strategies for

the future.

Hopefully, during the next two days:

1. We can continue to dispel myths. Myths, which can be

the greatest enemy of science; and

2. We can increase our knowledge and understanding of the

ROLE OF FIRE IN THE CENTRAL HARDWOODS.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 3

Abstract

The Central Hardwood Forest is an oak-dominated

deciduous forest that stretches from Massachusetts to

Arkansas and occurs in hilly to mountainous terrain. It is the

largest and most extensive temperate deciduous forest in the

world.

During the past 20 million years or so, angiosperms have

been gradually replacing gymnosperms as the dominant

plant form on earth, and deciduous hardwoods are

particularly adapted to the fluctuating seasonal climate and

moderate rainfall associated with the Central Hardwood

Region. As the glacial ice sheets retreated about 12-14

thousand years ago, forests re-invaded the region. Native

peoples populated the area and practiced extensive burning

to improve habitat for game and to aid in land clearing for

agriculture. By the time the early Europeans arrived, about

450 years ago, the Native People had adopted a sedentary

lifestyle and their population had increased to levels

comparable to that of Western Europe. The first European

explorers brought with them diseases that decimated the

native populations, and for almost 200 years the central

hardwood forest re-grew to become the “primeval forest” of

legend.

By the late 18th century, settlement by subsistence farmers

using European methods (draft animals and metal tools) was

in full swing. Much of the tillable land in the central hardwood

region was cleared, including substantial areas of steep and

hilly land. The industrial revolution in the post-Civil War

period ushered in a trend away from subsistence farming

and marginal land was abandoned to revert back to forest.

The last 100 years has seen an era of exploitive logging in

the central hardwood region, followed by a re-growing forest.

Human influences have predominated in shaping the present

forest and these include logging, burning, grazing, fire

control, wildlife management and pest introductions. In the

past 30 years, societal attitudes toward land and forests has

had a profound effect on governmental policy as it relates to

forestry. The maturing central hardwood forest, mostly

owned by private individuals, is rapidly becoming a “resource

at the crossroads”.

Introduction

The historical developments that shaped the central

hardwood region are significant in understanding how the

present forests came to be and, as always, knowledge of

history enables us to learn from past successes and failures.

Thus, history provides not only the background, but also the

context for future management. For purposes of this

discussion, the history of the central hardwoods will focus on

the period after the last ice age (the period of human

impact). Because the particular forest that exists today is

mostly a result of events that have occurred during the life of

the trees that presently comprise it, this period will be

emphasized.

The overriding factors that predispose a given geographic

region to a particular vegetation type are the climate and soil

(site), and this has been true throughout the hundreds of

millions of years that vegetation has existed on earth.

Furthermore the prevailing vegetation creates habitat for

animal forms which are adapted to it, and they, in turn, have

a biofeedback relationship with the vegetation. This type of

association is the basis for what is called an “ecosystem”

(Odum 1971).

Evolution and the Deciduous Forest

The ability of plants to capture solar energy by using readily

available elements (carbon, hydrogen, oxygen, etc.) has

been the key to all biological development on earth. Fossil

evidence is our window to the past, and although incomplete,

it appears that this process of photosynthesis first developed

in primitive marine algae between 400 and 500 million years

ago (Eyre 1963). From these evolved the primitive land

plants that occupied the earth’s surface prior to the

Carboniferous Period (more than 200 million years ago).

During the Carboniferous Period (215-280 million years ago),

ferns and their relatives dominated the earth. Around 190

million years ago, the gymnosperms began to take over as

the principal vegetation, and they have given way to the

dominance of angiosperms (including deciduous hardwoods)

during the last 50 or so million years. The angiosperms,

though relatively recent in their development, have shown a

remarkable ability to adapt to the changing conditions of the

earth. From this point of view, it might be said that the central

hardwoods represent the cutting edge of evolutionary

change, as compared to the boreal conifer forest.

Perhaps the most recent (and most significant) episode of

climatic and vegetational change to affect the distribution of

present-day vegetation was the so-called ice age. The

continental glaciers extended down through North America

into southern Illinois, and a zone of tundra extended well

down into much of the present central hardwood region

(Davis 1983). As the Wisconsin ice sheet began to retreat

northward about 14,000 years ago, a boreal forest type

replaced the tundra and vestiges of this forest still exist as

the high-elevation spruce and spruce-fir communities in the

Appalachians (Maxwell and Davis 1972). The deciduous

forest community has gradually become the dominant

vegetation of the region and now forms the most extensive

forest of this type in the world.

Humans And Fire: A History Of The Central Hardwoods1

Ray R. Hicks, Jr.2

1

This paper was previously published in the Proceedings of the

11th Central Hardwood Forest Conference, Columbia, MO. 1997.

2

Professor of Forestry, West Virginia University, Morgantown,

WV 26506-6125.
4 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Impact of the Native People on the

Pre-columbian Forest

As important as the ice age , and its retreat, has been, the

impact of humans has probably been just as significant in

shaping the central hardwood forest. The indigenous people

occupied the central hardwood region for at least 10,000

years before Europeans came (Lesser 1993). In the

southern portion of the region, human occupancy probably

dates back as long as 14,000 years ago owing to the more

hospitable climate farther south (Buxton and Crutchfield

1985). Archaeologists classify the human occupancy of

North America into Late Prehistoric, Mississippian,

Woodland, Archaic and Paleo periods (Lesser 1993). Table 1

shows the chronology of these periods and outlines their

cultural lifestyle as well as broad climatic and vegetational

environments (Buckner 1992). Prior to the Woodland period,

indigenous peoples were nomadic hunter-gatherers. Their

influence on the forest during this period was most likely

through the setting of fires. Accidental human-caused fires

no doubt occurred during this time, and perhaps intentional

fires were set to drive out game or to retain habitat.

MacCleery (1992) speculates that the early Europeans’

observations of abundant game (bison, elk, etc.) in the

central hardwood region indicates a greater proportion of

grassland occurred than currently exists, which implies that

burning probably took place. Such burning, most likely would

have been the work of the indigenous population.

Around 1,000-500 B.C., the Native Americans adopted a

more sedentary lifestyle, and at some point during that time,

they began the practice of agriculture (Swanton 1979). They

also constructed elaborate burial mounds which can be

found in places, such as Moundsville, West Virginia, Marietta

and Hillsboro, Ohio, East St. Louis, Illinois, and Cartersville,

Georgia. During this agricultural phase, which extended into

the Late Prehistoric Period until European contact (about

A.D. 1600), the Native Americans generally lived in

permanent villages, often on the level land of river

floodplains (Davis 1978). The village would often have a

structural layout with houses of thatch or pole construction

surrounded by gardens and fields. During this time, the

Native Americans used tools as well as fire to clear land.

MacCleery (1992) states the case as follows: “The south was

dominated by fire-created forests, such as long-leaf pine

savannas on the Coastal Plain and Piedmont. The hardwood

forests of the Appalachian Mountains were also burned

frequently by native peoples. Virginia’s Shenandoah Valley

— the area between the Blue Ridge Mountains and the

Alleghenies — was one vast grass prairie. Native peoples

burned the area annually.” MacCleery further states: “On the

western fringe of the eastern forest, fire-dominated forests,

such as oak and oak-pine savannas, covered tens of millions

of acres.” Martin and Houf (1993) indicated that remnants of

so-called “balds” still exist in the Ozarks of southwestern

Missouri. These grassy glades usually occur on dolomitic

limestone soils, and require periodic fires for their

maintenance. They tolerate intermittent grazing, as would

have occurred when migratory herds of bison occupied the

area, but under year-round grazing imposed by the

European settlers, eastern redcedar began to overtake the

glades. Beilman and Brenner (1951) use several lines of

evidence to indicate that extensive forest cover is a relatively

recent development in the Ozark region. They refer to

Houck’s (1908) account of Ferdinand DeSoto’s observations

of the region in 1541 where he described fertile alluvial

bottoms planted in maize, pecans, plums and mulberry trees.

The open, park-like countryside was in contrast to the

hardwood and pine forests that exist today. Another

description of park-like vegetation in the Ozark region was

provided by the explorer Coronado, where he noted the

hunting of buffalo by the Osage Indians. Although substantial

prairie-like areas probably occurred throughout the central

hardwood region, Steyermark (1959) indicates that even

within the Ozark region, where grasslands were common,

substantial areas of hardwood forests existed prior to

European settlement.

Because the indigenous people tended to settle on

floodplains of major rivers, their influence was probably

greatest in such areas. Therefore, in many of the regions

that are dominated by uplands where rivers have narrow

floodplains, such as interior West Virginia, western Kentucky,

southeastern Virginia and into the Cumberland Plateau of

Tennessee, the evidence of Native American influence is

more limited. However, their use of fire may have enabled

them to exert an influence far removed from their

settlements. Cronon (1983), with reference to the American

Indian influence on the landscape states: “It is tempting to

believe that when the Europeans arrived in the New World

they confronted Virgin Land, the Forest Primeval, a

wilderness which had existed for eons uninfluenced by

human hands. Nothing could be farther from the truth.—

Indians had lived on the continent for thousands of years,

and had to a significant extent modified its environment to

their purposes.” The population density of Native Americans

is still being debated, with estimates ranging from 1 million to

more recent higher estimates, up to 18 million (Buckner

1992). At higher levels, their impact on the forest would have

been greater, but in any event, they most certainly had a

significant effect on the central hardwood region.

Post European Settlement to 1860

The first Europeans in North America, apart from settlers in

colonial villages along the Atlantic seaboard, were essentially

hunter/trapper/explorers. They had a limited direct impact on

the forest, but their indirect impact was profound. They

brought very little with them in the way of equipment,

livestock or supplies, but they did bring something that

proved to be more significant than any of these—disease. As

Williams (1989) points out, the epidemic of disease

introduced by the Europeans affected the native peoples to

such a degree that between 1520 and 1700, there was a

return to a more forested landscape in North America

(Buckner 1992). It was this forest that most historians refer to

as the “impenetrable ancient forests.” Early explorers kept

few records of their observations, but they opened the way

for others including botanists, surveyors, etc. Britain offered

huge grants of land to individuals, such as Lord Fairfax and

William Penn, to encourage settlement of the new world and

to gain control of the land from the native population. By
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 5

1600 A.D. waves of European immigrants, looking for a

better life and an opportunity to escape the feudal system,

made their way to the new world.

Most early settlers were subsistence farmers, a technology

the Europeans had adopted 1,000 years earlier (Blethen and

Wood 1985). The European model had proved very

successful in their homeland, therefore, the immigrants set

about clearing the North American forest to plant their crops

and graze their livestock, in much the same way as they had

“tamed” the European landscape about 1,000 years earlier.

They brought with them two things the native people

lacked—metal tools (ploughs, axes) and draft animals. Their

attitude was that the forest represented an obstacle to be

conquered. The forest also harbored wolves, mountain lions

and bears which posed a threat to domesticated livestock,

such as sheep and hogs.

Although the forest was a challenge to be overcome, forests

also provided many of the early settlers’ needs. For example,

nearly every family lived in a wooden house and kept their

livestock in a wooden barn (usually log construction).

Table 1—Chronology of Native People from Buckner (1992)

Cultural Event Date Years Ago Climatic/Vegetational Stages

HISTORIC PERIOD

Modern Times. Settlement Times. 2000 0 Man-made forests widespread.

High Indian mortality. Exploitation of forests and soil.

America discovered. 1500 500 Indian impacts (cultivation and fire)

MISSISSIPPIAN PERIOD mold forest character.

Indian cultures largely agrarian,

large palisades. 1000 1000

WOODLAND PERIOD

Pottery 500 1500

Corn cultivated; bow and arrow

AD Northern pines had moved into Canada

Burial mounds 0 2000 while southern pines had moved into

BC Tennessee - their present distributions.

ARCHAIC PERIOD 1000 3000

Marked increase in Indian

population; exchange with

other regions. 2000 4000 Sea level rises to modern position.

Beginnings of cultivation with

fire as the only feasible tool

for land clearing. 3000 5000 “Southern pine rise” results in marked

increase in dominance of southern

pines in SE.

Archeological evidence that 4000 6000

Archaic Indians used total

landscape of So. Appalachians.

5000 7000 Increased summer warmth and drought.

6000 8000

7000 9000 Central hardwood oak-hickory forests

became established.

8000 10000 Periglacial climate extended as far south

as an east-west line thru Ashville, NC

PALEO-INDIAN PERIOD

Largely hunting/gathering tribes;

fire was an available tool. 9000 11000 Temperate, deciduous forests replace Jack

pine/spruce/fir/larch in TN and NC.

First evidence of humans in SE 10000 12000

15000 Jack pine, spruce and fir are the primary

forest types as far south as Tennessee.

18000 FULL GLACIAL MAXIMA
6 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Additionally, wood was the primary fuel for heating. It was

not uncommon for a family to use 20-40 cords of wood

annually to heat and cook (MacCleery 1992). Wood was

used for fencing. MacCleery (1992) estimated that it required

about 8,000 fence rails to enclose a 40-acre square field,

and by 1850 there were about 3.2 million miles of rail fence

in the United States (mostly in the East). Potash was another

important product from American forests in the early to mid

eighteenth century. The demand for potash in Britain was

great during that time. It was used as a soil amendment and

as an ingredient in industrial processing (Williams 1989).

Hardwoods, especially oaks and maples, produced the

highest amounts of potash per unit wood burned and were

preferred by potash producers.

From the middle 1700’s into the 1800’s, many naturalists

trekked across North America in pursuit of unknown,

unnamed or undescribed species of plants and animals.

Some were students of Carolus Linnaeus, the Swedish

botanist. Others, such as Andre Michaux (1805) and John

Bartram (1751) and his son William (1791) published

detailed accounts of their travels through the central

hardwood region. From their descriptions and others, such

as surveyors’ field notes, we can get an idea of what the

forest was like at that time.

By this time the Native American populations had been

decimated by disease, and the deciduous forests had been

virtually free from their effect for between 150 to 250 years.

These observers wrote about vast forests of hardwoods.

Stephenson (1993) quotes Diss Debar’s description of the

high-elevation forests of West Virginia on the “table-lands” of

the Cheat and Greenbriar mountains: “Here, also neither

Oak, Poplar nor Hickory are to be found, but in their room

thrive noble specimens of Sugar Maple, Ash, Beech, Birch,

Wild Cherry and Black Walnut.” But observers who wrote

about the Ridge and Valley and Appalachian Plateau forests

of lower elevation noticed an abundance of oaks, hickories,

maples and yellow-poplar. Using the frequency of mention as

an index to species abundance, from John Bartram’s (1751)

travels through Pensilvania [sic] to Onandaga, the following

ranking of species in order of importance was developed

(Table 2). Although the age and size of trees and extent of

the forest was greater in 1750 than it is today, the species

mix looks fairly typical of present-day stands in the northern

part of the central hardwood region.

The early immigrants settled along the Atlantic coast and

generally used rivers as a primary means of transportation.

Westward progress of settlement was relatively slow from

1600 well into the 1700’s, mostly due to the lack of a

transportation system. The Fall Line of the Peidmont marked

the limit of navigability on most eastern rivers; thus, many

settlements were established at this position along the

frontier. Innovative feats of engineering created water

transportation systems, such as the C and O Canal, brainchild

of George Washington, but it was only possible to proceed

from the Chesapeake Bay to Cumberland, Maryland. The

Allegheny Front proved too formidable for the technology of

the day. It was the railroads that finally opened the land

beyond the Alleghenies. Even though roads (such as the

Drover’s Road in North Carolina) were used during this time,

they were not suitable for transporting heavy materials.

Lewis (1995) indicated that it was the construction of the

railroads into West Virginia’s back counties that allowed a

“timber boom” to occur and transformed subsistence farming

into “commercial” agriculture in the state.

A fledgling iron industry was developing in the eastern United

States, especially in the Appalachian Plateaus. Low-grade

iron ore was processed in stone furnaces using local

limestone for purification and charcoal for heating. Charcoal

was produced by clearing patches of forest 1-3 acres in size

(Clatterbuck 1990). Luther (1977) estimated that the 11

furnaces operating in the mid-1800’s on the Highland Rim of

Middle Tennessee required 375 square miles (240,000

acres) of timber to support them. Similar operations in West

Virginia, Pennsylvania and elsewhere combined would have

consumed several million acres of forestland in the central

hardwood region.

The Industrialization Period, 1861-1929

As is often the case in a post-war era, the Reconstruction

period following the Civil War ushered in a period of

sweeping change. The steam engine, which could be moved

from place to place, was taking the place of water power for

milling. The factories of the North were being converted to

peacetime production. And the development of a massive rail

transportation system was well under way, spurred on

initially to supply troops in the war (Fig. 1). With the end of

slavery, many large plantation farms were unprofitable. Many

subsistence farmers throughout the central hardwood region

gave up farming for a more lucrative lifestyle to work in

factories, mines or logging camps. Extensive tracts of land

were purchased and consolidated by large timber or mineral

companies as a speculative enterprise (Eller 1985). The

migration of farmers to logging, milling or mining camps was

taking place nation wide (Fig. 2), but was more pronounced

in areas, such as the central hardwood region, where small

subsistence farming predominated.

In addition to the emergence of America as an industrial

nation, the Industrial Revolution brought about significant

changes in agriculture. Tractors powered by steam, and later

gasoline, replaced draft stock, and the use of fertilizers and

genetically improved plants and animals increased

production per unit area. Up until about 1908, as the

population of the United States grew, more land needed to

be placed under cultivation, and, conversely, land was taken

out of forest production (Fig. 3). The amount of forestland

and cropland has stabilized with the small additional

withdrawals of forestland coming from urbanization and

rights-of-way for highways, pipelines and power

transmission. Mechanization of agriculture also caused a

shift in the type of land that could be easily cultivated. Much

of the steeper land within the central hardwood region

became submarginal and was abandoned. The capital

investment required to buy large machinery put agriculture

on more of a business level than that of a homesteading

operation.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 7

Several secondary effects of the industrialization period on

the central hardwoods were also felt. For example, the

factories, machines and vehicles needed fuel which

promoted rapid development of the fossil fuel industry. Coal,

oil and gas production was spurred on throughout the central

hardwood region. Forests were withdrawn from production

by surface mining, roads, well sites, pipelines, etc. In

addition, the booming economy required wood for

construction of factories, new housing for factory workers

(either immigrants or relocated farmers), mine timbers, etc.

The booming economy ushered in by the industrial revolution

and the vast resource base in North America swept over the

timber industry as well. According to Frederick and Sedjo

(1991), production of forest products tripled in the United

States between 1860 and 1910 (Fig. 4), mostly as a result of

increased production of lumber. Lumber production in West

Virginia, where hardwoods predominated, showed a similar

trend (Fig. 5). The economic boom in North America for

consumer goods continued until the beginning of the Great

Depression, but the timber boom had peaked in the central

hardwood region by 1920, mostly as a result of overcutting.

Ahern (1928) in his booklet “Deforested America” stated the

situation as follows: “In 1919 the annual drain on our forest

resources was estimated at four times the annual growth.”

Other estimates, although less dramatic, show the same

trend where drain exceeded growth during this period (Fig. 6).

The history of the North American logging boom is the

subject of several books (Blackhurst 1954, Brown 1923,

Clarkson 1964, Fries 1951), and like similar events, such as

the California Gold Rush, the timber boom is surrounded by

a certain amount of folklore as well as fact. But, in any event,

it had a more dramatic effect on our present day central

hardwood forest than any single event. Almost all the forests

that had any merchantable value were cut over during that

period. Only small pockets remained, such as found in the

Joyce Kilmer Memorial Forest in North Carolina (Lorimer

1976). The use of steam engines for logging and steam

donkeys for skidding exacerbated the effect in causing

numerous fires which burned repeatedly through the logging

slash and forest floor, exposing the mineral soil to erosion.

Eller (1985) quotes Thomas Wolfe of Ashville, North

Carolina, as follows: “The great mountain slopes and forest

had been ruinously detimbered; the farm-soil on the hillsides

had eroded and washed down; high up, upon the hills one

saw raw scars of old mica pits, the dump heaps of deserted

mines. . . . It was evident that a huge compulsive greed had

been at work.” Boom towns, such as Davis and Spruce,

West Virginia, Tellico Plains and Gatlinburg, Tennessee, and

Fontana, North Carolina, developed rapidly; some are

completely gone today, and others turned to other industry or

tourism to survive. But in spite of the apparent devastation,

the forests of the central hardwood region started to regrow,

resulting from natural regeneration (stored seed, seed

distributed by wind or animals, seedling sprouts or stump

sprouts). This period of recovery is the focus of the following

section.

The Regrowing Forest, 1930-1996

Well before 1930, several persons in positions of leadership

in American forestry had expressed grave concern about the

state of the forest in North America. Gifford Pinchot,

Table 2.—Summary of frequency of mention by species (or species groups) from

Bartram’s (1751) trip through Pennsylvania

Species Number Times Mentioned Ranking

white oak and black oak 25 1

white pine 12 2

chestnut 10 3

spruce 10 3

hickory 8 4

sugar maple 8 4

linden 7 5

pitch pine 7 5

elm 6 6

beech 6 6

white walnut (butternut?) 6 6

birch (yellow?) 5 7

poplar (yellow-poplar?) 4 8

ash 4 8

sugar birch (black?) 3 9

great magnolia (cucumber?) 3 9

locust (black?) 2 10

walnut (black?) 1 11

hophornbeam 1 11

plane (sycamore?) 1 11
8 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Figure 1.—Clarkson’s (1964) map of West Virginia railroads in 1917 and band saw mill towns, 1875-1920.

Figure 2.—Farm and non-farm

population, 1880-1988, from

MacCleery (1992).

generally regarded as the father of American forestry, stated

his opinion in Ahern’s (1928) publication, as follows: “Forest

fires are steadily growing worse in America, and fire

prevention is absolutely indispensable. But the axe,

carelessly used is the mother of forest fires. The axe and not

fire is our greatest danger. Until the axe is controlled there

can be no solution of the fire problem, or of the problem of

forest devastation in America.” Pinchot, who was a

consummate politician, had been trained in forestry in

Europe. His first job had been as a forester on the huge

Vanderbilt estate near Ashville, North Carolina, in 1892 (later

to become the site of the Biltmore Forest School), but his

greatest accomplishments were in the political arena. He

became a Washington political insider and the first Chief of

the USDA, Forest Service. He was instrumental in

establishment of the Yale School of Forestry and the Society

of American Foresters. Pinchot’s influence on the central

hardwood forest came mostly through the establishment of
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 9

Figure 3.—Cropland vs forest area,

1850-1980, from MacCleery (1992).

Figure 4.—Domestic Production,

1800-1910.

the eastern National Forests. Earlier, Pinchot’s influence on

then President Theodore Roosevelt had resulted in

substantial additions to the national forest reserve between

1905-1909. About 80 million acres of federal land in the

western states had been set aside as forest reserves as

Pinchot and others became increasingly concerned about

overcutting in the East.

But the eastern lands had been purchased by private owners

and were not so easily annexed by the federal government.

The breakthrough came in 1911 with passage of the “Weeks

Act” which authorized the federal government to purchase

lands in eastern America. Initially, the purpose for acquiring

the land was to protect headwaters of navigable streams.

Lands which formed the nucleus of many of our current

National Forests were purchased under the Weeks Act

authority. Almost all the initial purchases were cut-over, and

often burned-over, lands belonging to large corporate owners

and were bought for prices of $2 to $4 per acre; hence, they

were called “the lands that no one wanted.”

In 1924, the Clarke-McNary Act added the production of

timber to the mandate for National Forest lands. By 1930,

the Great Depression was in full swing, and land prices,

especially the cut-over forest lands, were extremely low, as

corporate owners often viewed the land as a tax burden with

no foreseeable harvest of timber in sight. For example, in

1933 a purchase of almost 330,000 acres for the

Monongahela National Forest was made at an average cost

of $3.43 per acre (USDA, Forest Service 1970). From these

beginnings, the eastern National Forests developed (Fig. 7),

many within the central hardwood region.

The Weeks and Clark-McNary Acts, in addition to mandating

the acquisition of land, also provided funding to states for fire

control. The Forest Service also became the lead agency in

promoting forest fire control at the federal level. The results

came slowly at first, but by the 1960’s, forest fires which

consumed an average of 40-50 million acres in the 1930’s

were reduced to 2-5 million acres. (USDA, Forest Service

1987) (Fig. 8). Although the central hardwood region is not

as fire prone as conifer-dominated regions of the country, fire

has always been a significant disturbance in the oak forests

of the central hardwood region (Van Lear and Waldrop

1989). For example, in 1987 more than 50 percent of the

land area in Mingo County, West Virginia burned over (Hicks

and Mudrick 1993) (Fig. 9).
10 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

In 1933, President Franklin D. Roosevelt signed the

Civilian Conservation Corps bill into law. The CCC was

part of Roosevelt’s “New Deal,” intended to solve two

problems (unemployment and environmental

destruction). It was estimated that nearly 250,000

young men who were of employment age, were out of

work during the 1930’s and Major Stuart, Chief of the

USDA, Forest Service, suggested that the army be

used to build camps and generally administer the

program. But unemployed civilians made up the work

force. Many CCC camps were established within

National Forests and their activities included planting of

trees, fire control, watershed improvements, road

improvement and construction of recreational facilities.

Although their impact on today’s central hardwood

forests are largely undetectable, they planted millions

of trees (mostly conifers) on denuded lands, and many

of the recreational structures they built are still usable

today, almost 60 years later.

The Depression drove more people from marginal

farms, and the trend in reversion of farmland to forest

continued through the 1930’s. In 1942, the Depression

ended with America’s entry into World War II. The war

accomplished what governmental programs could not by

employing millions of Americans, either as soldiers or in

the factories that manufactured war materials. The war

put technological development into high gear, and many

of the devices that have significantly affected our society

were developed during this period. The post-war

economy was booming when soldiers returned home,

many to take factory jobs, marry and raise families. The

impact of these developments on the central hardwood

region was a shift in the lifestyle and objectives of land

owners. Many people kept their residence in a rural

setting while seeking employment in mills and factories

elsewhere. The reason for owning land shifted away from

utilitarian (production) objectives toward recreational/

aesthetic enjoyment/ residential objectives (Birch and

Kingsley 1978) (Fig. 10).

The changing land ownership characteristics have

continued into the most recent decades throughout the

central hardwood region. This is illustrated by the changes

that occurred from 1957 to 1975 in West Virginia, a state that

experienced little or no total population growth over that

period. In 1957, the West Virginia Forest Industries

Committee reported that there were 133,570 private owners

of forestland, who owned over 8.9 million acres. The average

size of ownership was 66.6 acres in 1957. By 1975, the

number of owners had increased to 207,500 and the

forestland acres had also increased to 10.3 million acres. But

the average size of ownership had dropped to 49.8 acres

(Birch and Kingsley 1978). Twenty percent of the land

owners in 1975 were not from rural or farm backgrounds.

Twenty-two percent of the land owners held their land

primarily for aesthetic enjoyment or recreation, and another

23 percent owned the land primarily for their residence. Only

3 percent owned land primarily for timber production. Similar

changes have taken place throughout the central hardwood

region.

Alig (1990) reported the trends in forested land by states. If

the 11 states that constitute the bulk of the central hardwood

region are summarized (Table 3), the trend in total forestland

is depicted in Figure 11. A decrease of about 2.5 percent in

forestland is projected to occur by 2040, mostly due to urban

expansion.

Agricultural policy continued to have an impact on forestry

throughout the post-war period. The Conservation Reserve

Program in the 1960’s provided compensation to farmers to

convert cropland to forest by planting trees. The Forest

Incentives Program provide cost sharing to small private land

owners for tree planting and other forest practices. The

Stewardship Incentives Program of the 1990’s provides

federal cost sharing to small forest land owners to develop

and carry out planned forest management. Some states

have offered additional property tax incentives to land

owners enrolled in the Stewardship Incentives Program.

Figure 6.—Timber growth/removals from MacCleery (1992).

Figure 5.—West Virginia lumber production from Zinn

and Jones (1984).
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 11

These programs are designed to promote good forest

management among the many and varied small private land

owners, a type of ownership that predominates in the central

hardwood region. Technical support for forestry is provided

to private land owners through state forestry agencies,

Agriculture Extension Agents and the USDA Forest Service,

State and Private Forestry. But in spite of this large effort, the

level of participation among private forest land owners in the

central hardwood region has generally lagged behind

government expectations, partly due to the inherent

independence of property owners, partly due to a lack of

interest or economic incentives and partly due to an inability

to inform land owners of the availability of such programs.

The withdrawal of forestland for agriculture leveled off in

about 1933, but during the post-war decades, some decline

in forest area has occurred. This has mostly resulted from

development-related activities. One example is the interstate

highway system which was developed during the 1960’s and

1970’s for the most part. Rights-of-way for gas transmission

and electric power transmission also consume large

corridors through the forest, and surface mining and urban

development have taken up significant amounts of

forestland. Topography and economic factors have resulted

in the central hardwood region’s being an area that is

currently among the highest in the eastern United States for

coverage with natural vegetation (Kolpatek and others 1979)

(Fig. 12).

Some of the most significant changes to the central

hardwood forest in recent years have been brought about as

a result of human introduction of insects and disease

organisms. These changes are often not dramatic, but their

impact is unquestionably large. The list of important

introduced pests includes dogwood anthracnose, Dutch elm

disease, oak wilt and beech bark disease. But the two with

the most impact or potential impact on the central hardwood

forest are chestnut blight and gypsy moth. The former is a

disease that has effectively eliminated a species, which at

one time was among the most widely distributed and

economically important in the central hardwood region. The

latter is an introduced insect that continues to spread

Figure 7.—Distribution of National Forests.

Figure 8.—US wildfire trends,

from MacCleery (1992).
12 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

southwestward through the central hardwood region and

seems to have the potential to spread through the entire

region over the next 50 to 75 years.

The chestnut blight fungus (Cryphonectria parasitica)

was introduced to the New York Botanical Gardens in

1904 (Giddings 1912, Murrill 1904). By 1915 most states

east of the Mississippi River reported infected trees. By

the late 1930’s, most mature chestnut trees were dead

throughout the eastern states. The following should

provide a frame of reference for the significance of

American chestnut. Chestnuts were a source of food and

commercial enterprise for mountain farmers. During the

settlement period, log cabins were often constructed of

chestnut logs due to their durability and decay

resistance. Most split-fence rails and posts were

constructed of chestnut where it was available. And due

to its durability, chestnut lumber was widely used in

construction of barns, sheds and even for the

manufacture of furniture. The bark of chestnut was an

important source of tannin for leather tanning factories.

Chestnut also occupied an important position in the

hardwood ecosystem. Braun (1950) used the name

“oak-chestnut” to identify one of the most significant

forest formations in the eastern deciduous forest.

American chestnut was an important source of mast for

a variety of wildlife species (animals and birds) most of

which served as prey for larger predatory mammals.

The impact of the loss of American chestnut can never

be fully assessed. But Brooks (1915) indicated the

following facts regarding chestnut in West Virginia

alone. The annual harvest of chestnut in West Virginia

in 1915 was 118 million board feet. The entire volume

of chestnut in West Virginia was estimated to be 5

billion board feet. At a stumpage value of $8 per

thousand board feet, in 1915, the entire chestnut

inventory was estimated to be worth $55 million. At

current stumpage prices for red oak that value would

be about $1.5 billion. The only evidence remaining of

the once important chestnut are occasional root

sprouts, which usually survive to the sapling stage

before being killed by the blight, and the grey remains

of stumps of salvaged trees. Ironically, many of these

stumps still have coatings of charcoal, evidence of the

fires that burned over the area, probably after the

chestnuts had already succumbed to the blight.

Gypsy moth (Lymantria dispar) is an introduced insect

that potentially could have as great, or greater, impact

on the central hardwoods than chestnut blight. Gypsy

moth was inadvertently introduced into the Boston area in

1869 by Leopold Trouvelot (Liebhold 1989), who was

interested in hybridizing the gypsy moth with native

silkworms. The spread of gypsy moth has continued, and it is

potentially capable of becoming established throughout the

central hardwood region. Although gypsy moth can defoliate

a variety of tree species, including conifers, deciduous

species are their preferred hosts. Oaks are among the most

preferred hosts of gypsy moth (Bess and others 1947,

Gansner and Herrick 1987). Black cherry, hickory and

maples are intermediate in preference while species, such

as yellow-poplar and ashes, are virtually resistant. The

significance of this to the central hardwood region is the fact

that susceptible species make up the bulk of the forest

composition throughout the region, the exception being

yellow-poplar, especially in the mixed mesophytic forest.

Figure 9.—Percentage of West Virginia counties burned in 1986,

1987 and 1991.

Figure 10.—Reason for owning land in West Virginia

(Birch and Kingsley, 1978).
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 13

Currently, gypsy moth has become established in, and is

endemic to, an area stretching from Michigan to North

Carolina (Fig. 13). But isolated populations occur throughout

the central hardwood region, having been spread by

movement of vehicles, logs or other objects containing egg

masses.

As a defoliator, gypsy moth’s impact on the tree is to reduce

its photosynthetic surface. The major defoliation occurs

during the spring (May-June) which is a period of critical

importance to deciduous trees since the soil moisture is

generally high and the temperature is moderate during this

time. Usually one defoliation is not lethal to trees, but a

single defoliation during a drought year or multiple years of

defoliation can lead to extensive tree mortality (Fosbroke and

Hicks 1989). The factors that precipitate extensive gypsy

moth outbreaks include presence of a suitable host plus

climatic conditions that favor buildup. Since gypsy moth is

situated in an oak-dominated region, the host is generally

suitable, thus outbreaks can occur whenever weather

triggers them. As can be seen in Figure 14, cycles of

outbreaks seem to occur about every 10 years. The

outbreaks in recent years have been affecting more acres

than in previous outbreaks, mostly due to the spread of

gypsy moth to occupy a larger endemic range.

The gypsy moth story is still unfolding, but its impact on the

central hardwood forest has already been experienced

throughout the portion of the region where it has spread. As

reported by Gansner and Herrick (1987), oak experienced an

average mortality rate of 24.2 percent between 1979 and

1984. Quimby (1987) reported a similar result where summer

droughts plus gypsy moth defoliation in 1980-1983 resulted

in the premature loss of 68 million trees in Pennsylvania, a

mortality rate of 27.6 percent. Much of the mortality occurred

in oak stands. Hicks and Fosbroke (1987) reported similar

rates of tree mortality in southwestern Pennsylvania and

western Maryland. The forest invaded by gypsy moth as it

entered northeastern Pennsylvania has been dubbed “the

new frontier” by Herrick and Gansner (1988), and essentially

this new frontier represents the oak-dominated forest of the

central hardwood region. The reduction in oak stocking that

has occurred in the part of the central hardwood forest where

gypsy moth has become established will most likely

represent what can happen throughout the entire region.

Another episode in recent history that has had a significant

impact on the central hardwood forest is the decline and later

reestablishment of wildlife populations across the region. In

addition to white-tailed deer, elk and bison were found

extensively throughout the region prior to European

settlement. Those species were, in part, hunted into

extinction but mostly were displaced due to habitat

destruction by land clearing for farming. Farmers also

eradicated predator species, such as eastern mountain lion,

eastern timber wolf and black bear, since these predators

were a threat to their livestock. By 1910, bison, elk, mountain

lion and wolf had been eradicated from the central hardwood

forest, and white-tailed deer and black bear were found only

in small isolated populations in remote and mountainous

regions.

Wildlife advocates and hunters pushed for, and obtained,

stricter hunting regulations, such as seasonal hunting, bag

limits and bucks-only hunting. The results for white-tailed

deer have been remarkable. In the absence of natural

predators, and with the second-growth hardwood forest

providing prime habitat, the deer herd has risen sharply

throughout the central hardwood region. In fact, deer have

generally become a serious problem to forest regeneration in

Table 3.—Summary of forested lands by state in the central hardwood region

(Millions Acres)

Projected

States 1952 1962 1970 1977 1987 2000 2010 2020 2030 2040

NC 19.6 20.0 20.1 19.4 18.4 17.7 17.6 17.5 17.5 17.5

VA 15.5 15.7 15.9 15.9 15.4 15.0 15.0 15.0 15.0 15.0

AK 19.6 19.9 18.0 16.8 16.6 15.6 15.4 15.1 15.1 15.1

KY 11.5 11.6 11.8 11.9 11.9 12.1 12.2 12.3 12.5 12.5

TN 12.5 13.4 12.8 12.9 12.6 12.4 12.3 12.3 12.3 12.3

MD 2.9 2.8 2.7 2.5 2.5 2.4 2.3 2.2 2.1 2.1

MA 3.3 3.0 2.8 2.8 3.0 3.0 2.9 2.8 2.7 2.6

PA 14.6 16.3 16.1 15.9 16.2 16.3 16.3 16.0 16.0 15.8

WV 10.3 11.4 11.4 11.5 11.8 11.8 11.8 11.8 11.8 11.8

OH 5.4 6.0 6.4 6.9 7.1 7.3 7.3 7.1 7.0 7.0

MD 14.3 13.5 12.5 12.3 12.0 12.0 12.0 11.9 11.9 12.0

Total 129.5 133.6 130.5 128.8 127.5 125.6 125.1 124.0 123.9 123.7

Change (from 1987) +2.0 +6.1 +3.0 +1.3 0 -1.9 -2.4 -3.5 -3.6 -3.8

Percent Change +1.6% +4.8% +2.3% +1.0% 0 -1.5% -1.9% -2.7% -2.8% -2.9%
14 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

many areas. Deer consume 1-1.5 kg of vegetation daily,

consisting of twigs, acorns and some grass and herbs

(Smith 1993). Deer are selective and prefer seedlings of

oaks and maples to those of black cherry (Marquis 1981).

Overbrowsing by deer can result in development of a ferndominated understory where tree seedlings have a difficult

time getting established due to competition and allelopathy

(plant toxins). Thus, the long-term effect of deer on the

forest may be to retard the process of regeneration,

selectively change the species composition and reduce the

overall diversity of species.

The modern chapter in forest history also includes a marked

change in the social context of forest resources. This has

been brought about by a combination of factors including the

changing demographics in America, where people today are

physically removed from the land. But other factors have

come into play. For example, abusive and exploitive land

uses, such as exploitive timbering, unregulated mineral

extraction, air and water pollution and uncontrolled

development are all sources of public concern. The

environmental movement began to take shape as a cultural

phenomenon during the late 1960’s. Rachel Carson’s book,

Silent Spring, did for environmentalism what Harriett

Beecher Stowe’s Uncle Tom’s Cabin did for the

emancipation movement. Against a backdrop of Kennedy

altruism and polarizing issues, such as the cold war, racism

and a budding Viet Nam conflict, it seems, in retrospect,

inevitable that the environmental movement, led by altruistic

urban youth, should flourish. Most foresters, at that time,

were from rural backgrounds and had come through their

forestry training with a paternalistic approach toward

resource management, being one in which the forester is

regarded as the trained professional, and should be trusted

to make the appropriate decisions regarding management of

the resource. Thus, the elements of conflict were in place.

The political arena became the venue for the struggle.

Environmental activist groups gained strength. The

Wilderness Act of 1964 represented a milestone in

environmental policy legislation whereby areas in the

National Forests could be set aside for “wilderness,” and

timber cutting in these areas would be prohibited (Dana and

Fairfax 1980). Controversy over the use of clearcutting on

public land resulted in the landmark Monongahela Decision

of 1970 that recommended uneven-age management as a

primary management policy for the Monongahela National

Forest in West Virginia. This decision was more significant

as a precedent than it was to the Monongahela National

Forest. In effect, it imposed the will of the public over that of

foresters, who had always been regarded as the experts in

forest resource management.

The issue of the 90’s seems to revolve around regulation of

private land. The so-called “spotted owl” controversy in the

Pacific Northwest has set the tone. The perceived situation

was one in which an endangered organism’s habitat (oldgrowth forest) was not limited to a single land owner but

extended to private land as well as public land.

The debate extending to small non-industrial private forest

(NIPF) owners has progressed from the passage of

regulations requiring the use of “best management practices”

for logging to a more comprehensive and controversial

debate over whether or not land use should be controlled at

the landscape level. If such control is needed to facilitate

what is called ecosystem management, how can it work in

areas, such as the central hardwood region, where over 75

percent of the land is owned by NIPF owners? Proposed

solutions include a combination of government educational

and assistance programs plus incentive programs (Campbell

and Kittredge 1996). The use of legislative regulation to

achieve the goals of ecosystem management is a

controversial approach (Argow 1996) and one that evokes

strong emotions. The fear among private owners- that they

will lose prerogatives to use their land- is not unfounded.

Examples abound where the “general good” of the public has

taken precedence over the desires of the individual.

On the other side of the issue are those representing private

land owners (Farm Bureau, National Woodland Owners

Association, etc.) who contend that the loss of management

options to a private land owner due to regulation constitutes

a “taking” under the Fifth Amendment of the Constitution, for

which the owner is entitled to be compensated. As these

debates continue, the outcomes will most certainly affect the

way forests can be, and will be, managed and exploited in

the central hardwood region in the years to come.

In summary, the history of the central hardwood forests is

one that is based on the actions and interactions of climatic

and biological factors, and, in more recent times it has been

strongly affected by humans and their actions (Table 1).

Figure 11.—Change in forestland (actual and predicted) for 11

states with significant area within the central hardwood region

(AR, KY, MA, MD, MO, NC, OH, PA, TN, VA WV), 1950-2040,

from Alig (1990).
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 15

80 %

60 %

40 %

20 %

Going back to the retreat of the last glaciers (12,000-14,000

years ago) and the climatic warming trend that has followed,

the deciduous forests found an ideal environment and

flourished in the central hardwood region. Indigenous people

used fire and cultivated land, and their activities peaked

about the time European settlement began. The popular

notion of an undisturbed “primeval” forest that predated

European settlement seems largely to be a myth. After native

people were decimated by disease, forests regrew for about

200 years, and then clearing for subsistence farming began

in ernest. This proceeded until about the time of the Civil War

and the onset of the industrial revolution. This period marked

the beginning of a shift in agriculture to more level land, and

abandoned hill lands in the central hardwood region began

to regrow into forest. A period of exploitive logging around

the turn of the 20th century has been followed by a

regrowing forest. Forest land owners and the public have

changed in their attitudes concerning forest resources in

recent times.

Now with many hardwood stands in the central hardwood

region approaching 100 years of age, the resource is

maturing. Many forces are mustering for a piece of the

action. Developers, speculators, industries, agencies,

advocacy groups and environmentalists are all becoming

aware of this unique and diverse forest. Our actions today

will be the history of the future, and foresters should be

prepared to participate in, and hopefully guide, the decisionmaking process at this important crossroad in the central

hardwood regions development.

Figure 12.—Coverage of natural vegetation by county in the US (Klopatek and others 1979).
16 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Figure 13.—Gypsy moth quarantine

area as of 1996.

Figure 14.—Outbreak history of

gypsy moth in the Northeast.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 17

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Pudup, M.B., Billings, D.B. and Waller A.L. University of

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of our problem. Gypsy Moth News. U.S. Department of

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Allegheny Mountains in the States of the Ohio,

Kentucky and Tennessee in the Year of 1802. London:

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Quimby, J. 1987. Impact of gypsy moth defoliation on

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Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 19

Abstract

It is now widely acknowledged that frequent low-intensity

fires once structured many western forests. What is not

generally recognized, however, is that most of those fires

were purposefully set by native people, not started by

lightning. Data from the Rocky Mountains attest to the

widespread use of fire by native people, as does the ecology

of aspen, the only common deciduous hardwood in the

West. Fire history studies all show that aspen once burned at

frequent intervals, yet aspen will readily burn only when the

trees are leafless and the understory dry — conditions which

occur only early in the spring before leaf-out and understory

regrowth, or late in the fall after leaf-drop and the understory

has been killed by frost. During both these periods, though,

there are few lightning strikes and virtually no lightningstarted fires in the Rocky Mountains. Thus if aspen burned

frequently in the past, as all evidence indicates it once did,

then those fires must have been started by native people,

who used fire to modify plant communities for human benefit.

Similarly, eastern deciduous forests will readily burn only

when leafless, but during that time there are few lightning

strikes — one of many indications that aboriginal burning

was also common in the eastern U.S.

Introduction

It is now generally acknowledged that frequent low-intensity

fires were once common and historically structured many

western forests (Barrett and Arno 1982, Kay 1995, Kloor

2000). In addition, it is widely assumed that most, if not all, of

those earlier fires were started by lightning (Kloor 2000,

Loope and Gruell 1973:434, Romme and Despain 1989).

Data from Arizona, New Mexico, Idaho, Yellowstone, and the

southern Canadian Rockies, though, all attest to the

widespread use of fire by native people, as does the ecology

of aspen (Populus tremuloides), the only common deciduous

hardwood in the West.

Aboriginal Burning in the West

Selway-Bitterroot Wilderness Area

Brown and Bradshaw (1994) and Brown and others (1994,

1995), for instance, compared the U.S. Forest Service’s

Prescribed Natural Fire Program with pre-European

settlement fires in the Selway-Bitterroot Wilderness Area

along the Montana-Idaho border. Based on stand-age

analyses and fire history maps, Brown and others (1994,

1995) determined how frequently various forest types burned

in the past and then compared those data with how

frequently the same vegetation types burned from 1979-

1990 when lightning-caused fires were allowed to run their

course. Brown and others (1994, 1995) reported that, on

average, the area burned during pre-European times was

nearly twice as great as the area burned by lightning fires

alone today. Moreover, low-elevation montane areas that

once had the highest fire frequency, now seldom burn. Since

the overall climate has not changed, it is unlikely that

lightning-caused fires burn less area today then they did in

the past. Instead, it is likely that there are fewer fires today

because native people no longer use fire to manage the

land, as they once did.

Southern Canadian Rockies

A similar situation exists along the east slope of the southern

Canadian Rockies. In the past, fires were exceedingly

frequent, while today lightning-caused fires seldom occur

(Barrett 1996, Heathcott 1999, Kay and others 1999, Kay

and White 1995, White 1985). In some vegetation types, firereturn intervals are now 100 times greater than they were in

the past. Lower montane valleys that once burned every 5

years or less now do not burn at all. Based on this and other

evidence, Parks Canada has concluded that native burning,

not lightning-caused fires, was critical in maintaining what

heretofore was believed to be the “natural” vegetation

mosaic of the southern Canadian Rockies (White and others

1998). That is to say, there simply are not enough lightningcaused fires to account for historical burn and vegetation

patterns (Heathcott 1999).

Yellowstone National Park

Prior to park establishment, Yellowstone’s northern

grasslands had a fire-return interval of once every 25 years

(Houston 1973). Yellowstone has had a “let burn” policy for

nearly 30 years now, yet during that period, lightning-caused

fires have burned practically none of the northern range. In

1988, fire did burn approximately one-third of the area, but

according to agency definitions that was “unnatural” because

the fire was started by a man, not lightning. Moreover, the

fire that burned the largest portion of the northern range

started outside the park and was driven by unusually strong

winds more than 70 km before it reached the northern range.

Besides, the 1988 fires are thought to be a 100-300 year

event (Schullery 1989a, 1989b), so similar fires could not

have caused the original 25 year fire frequency.

Despite a series of recent droughts, why has Yellowstone’s

northern range remained virtually unburnt? Park biologists

contend that this is because “lightning has chosen not to

strike very often on the northern range” (Despain and others

1986:109). That assertion, though, is not supported by data

from the Bureau of Land Management’s Automatic Lightning

Strike Detection System which shows that, on average,

lightning strikes the northern range 4 times per km2

/yr (Kay

1990:136-137). So lightning strikes, but why doesn’t the

Native Burning in Western North America:

Implications for Hardwood Forest Management

Charles E. Kay1

1Adjunct Assistant Professor, Department of Political Science,

Utah State University, Logan, UT 84322.
20 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

range burn? The answer is that when most lightning strikes

occur, the herbaceous vegetation is too green to carry a fire

(Kay 1995). Thus, it is likely that the park’s original 25 year

fire frequency was entirely the product of aboriginal burning.

Arizona and New Mexico

At European contact, ponderosa pine (Pinus ponderosa)

forests in Arizona, New Mexico, and throughout the Rocky

Mountains were open and park-like (see Figure 1), but have

since developed into impregnable thickets due to the

ingrowth of smaller trees, which, in turn, has created the

current forest health crisis (Covington and Moore 1994, Fule

and others 1997). The open nature of the original forests, as

well as the more recent proliferation of smaller trees, is

generally attributed to modern fire suppression and the lack

of lightning fires. That is to say, it is commonly believed that

lightning historically was the primary ignition source, not

native people (Seklecki and others 1996, Swetnam and

Baison 1996a). This interpretation, though, is not supported

by lightning frequency data or time of fire-scar analyses.

In the southwest, over 95% of lightning strikes occur after

July 1st (see Figure 2a), while, historically, 85% or more of

ponderosa pines were scarred by fire during April, May, and

June (see Figure 2b). Now, despite the relatively low

incidence of lightning, lightning fires early in the year do burn

a disproportional area due to generally dry conditions at that

time of year (Baison and Swetnam 1990:1562, Barrows

1978, Swetnam and Betancourt 1990), but lightning fires

alone still cannot account for the magnitude of early-season

fire scarring seen during pre-European times (Barrows

1978). In many mountain ranges today, there simply are not

enough lightning fires to have caused the high fire frequency

observed prior to European settlement (Baison and

Swetnam 1997:3). Thus, it is logical to assume that a large

proportion of the “natural” fire regime in pine forests and

Figure 1—An example of the ponderosa pine communities that were common throughout the West at European

contact. These early forests were so open and park-like that it was possible to go nearly everywhere with horsedrawn carriages. These pine forests also supported a lush understory of grasses and forbs, while today, forest

ingrowth has created a forest health crisis and understory forage production has fallen to virtually nothing (Uresk

and Severson 1989, 1998). Photo taken on the south rim of Arizona’s Grand Canyon by R. Arnold (No. 166) in

1905 — courtesy U.S. Geological Survey Photographic Archives, Denver, CO.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 21

Figure 2—(a) Frequency distribution of cloud to

ground lightning strikes in Arizona and eastern new

Mexico during 1989 and 1990. From Watson and

others (1994:1720). (b) Frequency distribution of firescar data for the southern Rockies. Clearly most

trees were scarred by fire when there were few

lightning strikes. Timing of fire scars determine by

microscopic analysis of when individual growth rings

were damaged by fire. From Brown and Sieg (1996),

Fule and Covington (1999), Fule and others (1997),

Morino (1996), Swetnam and Baison (1996b).

other regions of the southwest was actually due to aboriginal

burning (Bonnicksen 2000).

Aspen Ecology

Repeat photographs and fire history studies indicate that

Rocky Mountain aspen communities burned frequently in the

past, yet experience has proven that aspen is extremely

difficult to burn (Brown and Simmerman 1986). Terms such

as “asbestos type” and “firebreak” are often used to describe

aspen (DeByle 1987:75). Even raging crown fires in

coniferous forests seldom burn adjacent aspen communities

(Fechner and Barrows 1976). DeByle and others (1987:75)

noted that “wild fires that had burned thousands of acres of

shrubland or conifer types during extreme burning conditions

usually penetrated less than 100 feet into pure aspen

stands.” Lightning-fire ignition rates for aspen are also the

lowest of any western forest type, and overall ignition rates

are less than half that for all other cover types, including

grasslands (Fechner and Barrows 1976). At current rates of

burning, “it would require about 12,000 years to burn the

entire aspen type in the West” (DeByle and others 1987:73).

Something is clearly different today than it was in the past.

Research has shown that these communities will readily

burn only when aspen is leafless and the understory plants

are dry, conditions that occur only during early spring and

late in the fall (Brown and Simmerman 1986). Prior to May

15th and after October 1st, though, there are few lightning

strikes and virtually no lightning fires in the northern or

southern Rocky Mountains (Kay 1997a, 1997b; Nash and

Johnson 1993) — see Figures 3 and 4. So if aspen burned

at frequent intervals in the past, as fire-frequency data and

historical photographs indicate it did, then the only logical

conclusion is that those fires had to have been set by Native

Americans (Kay 1997a, 1997b).

Why did Native People Burn?

Thus, it is increasingly clear that Native Americans had a

major impact on ecosystems by repeatedly firing the

vegetation (Bonnicksen 2000). They did this to modify plant

and animal communities for human benefit and to increase

productivity (Pyne 1995). In California, for instance, native

peoples had 70 reasons for burning (Lewis 1973), and even

in northern Canada, where the vegetation is less diverse,

Native Americans still set fires for at least 17 different

reasons (Lewis and Ferguson 1988). Native peoples

commonly set fires to enhance forage production and to

attract prey; to herd wildlife during hunts; to rid the forests of

underbrush and to facilitate travel; to enhance plant

production such as berry producing shrubs, mast species,

and root crops; to destroy poisonous snakes and other

vermin; to clear the land prior to farming; to kill trees for

firewood; to rid the land of places an enemy could hide; and

as an instrument of war, among many others. Although
22 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

aboriginal burning has been widely reported in the

anthropological literature (e.g., Boyd 1986; Gottesfeld 1994;

Lewis 1985; Pyne 1993, 1995; Turner 1991), those findings

have largely been ignored by ecologists and modern land

mangers (Bonnicksen 2000, Kay 1995).

Determining how fires started, though, is critical because

“fires set by hunter-gatherers differ from [lightning] fires in

terms of seasonality, frequency, intensity, and ignition

patterns” (Lewis 1985:75). The majority of aboriginal fires

were set in the spring, between snowmelt and vegetation

greenup, or late in the fall when burning conditions were not

severe (Pyne 1995). Unlike lightning fires, which tend to be

infrequent and of high intensity, native burning produced a

high frequency of low-intensity fires. Aboriginal burning and

lightning fires created different vegetation mosaics, and in

many instances, entirely different plant communities

(Blackburn and Anderson 1993, Bonnicksen 2000).

Moreover, aboriginal burning reduced or eliminated the

number of high intensity, lightning-generated fires (Pyne

1993, 1995; Reid 1987). Once aboriginal fires opened up the

vegetation, then subsequent lightning fires behaved like

those set by Native Americans (Pyne 1993, 1995).

Aboriginal Burning in the East

There are several lines of evidence which indicate that

aboriginal burning was also common in eastern deciduous

forests (Bonnicksen 2000, Hamel and Buckner 1998). These

include the lack of lightning fire, the original structure of the

forests, and species composition changes that have

occurred since European settlement, among others.

Figure 3—Frequency distribution

of lightning strikes on the Fishlake National Forest in

southcentral Utah. When aspen is normally dry

enough to burn during early spring and late in the fall,

there are few lightning strikes. This is true throughout

western North America (Kay 1997a, 1997b; Kay and

others 1999). Lightning data (1985-1994) from the

Bureau of Land Management’s Automatic Lightning

Strike Detection System, Boise, ID as provided by the

Fishlake National Forest, Richfield, UT.

Figure 4—Frequency distribution of lightning-caused

fires on the Fishlake National Forest in southcentral

Utah. When aspen is normally dry enough to burn

during early spring and late in the fall, there are few

lightning strikes (Figure 3), and few lightning-started

fires. Although, there are virtually no lightning fires

capable of burning aspen, historical photographs

and fire-history data indicate that aspen burn

frequently in the past (Bartos and Campbell 1998).

This suggests that the earlier fires had to have been

set by native people. Forest fire data (1960-1996)

from the Fishlake National Forest, Richfield, UT.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 23

Lightning Fires

Although lightning is common in most eastern forests,

lightning-started fires are rare (Barden and Woods 1974,

1976; Bratton and Meier 1998; Harmon 1982; McCarthy

1923). This occurs because when lightning strikes are most

frequent during July and August, eastern deciduous forests

are too green to burn. Like western aspen communities,

eastern deciduous forests will readily burn only when the

trees are leafless and the understories dry — conditions that

generally occur only early in the spring or late in the fall, and

during both those periods there are few lightning strikes and

even fewer lightning-caused fires. Fire history studies,

however, have shown that prior to European settlement, fires

were common in the eastern U.S. — many more than can be

accounted for by lightning alone (Bonnicksen 2000:259-269,

Bratton and Meier 1998). Therefore, the only logical

conclusion is that burning by native people was once

widespread in many eastern forests, similar to conditions in

the West.

Forest Structure

At European contact, many eastern forests were open and

park-like, with little undergrowth (Bonnicksen 2000, Day

1953, Olsen 1996). Like ponderosa pine forests in the West,

most eastern forests were once composed of large, widelyspaced trees “so free of underbrush that one could drive a

horse and carriage through the woods” (Botkin 1990:51).

Like western forests, though, most eastern deciduous forests

are now choked with dense underbrush and smaller

regenerating trees. The only way to create open park-like

stands in either western or eastern forests is for those areas

to have been subjected to a high frequency of low-intensity

surface fires. In eastern deciduous forests this would have

required fires either early in the year before the trees leafedout or in late autumn after leaf-fall. During both those

periods, though, lightning-caused fires seldom occur. Thus,

the only way for eastern forests to have displayed the openstand characteristics that were common at European

settlement is if those communities had regularly been burned

by native people as part of aboriginal land management

activities.

Species Composition Changes

An even more compelling piece of evidence is the species

composition changes that have occurred in eastern forests

since European colonization (Bonnicksen 2000). For the last

3,000-4,000 years, or longer, much of the eastern United

States was dominated by oak (Quercus spp.), American

chestnut (Castanea dentata) and pines (Pinus spp.), all firetolerant, early successional species (Bonnicksen 2000;

Bratton and Meier 1998; Clark and Royall 1995; Cowell

1995, 1998; Delcourt and Delcourt 1997, 1998, 2000;

Delcourt and others 1986, 1998; Hamel and Buckner 1998;

Meyers and Peroni 1983; Olson 1996). Since European

contact, however, oaks and pines have been replaced by

late-successional, fire-sensitive species, such as maples

(Acer spp.) (Abrams 1998, Bonnicksen 2000, Botkin

1990:51-71). That is to say, the species composition of many

eastern forests had been maintained for thousands of years

by frequent fires — fires, as we have seen, which could only

have been set by native people. It is equally clear that

aboriginal burning created the many eastern prairies and

“barrens” reported by early Europeans (Barden 1997, Belue

1996, Bonnicksen 2000, Campbell and others 1991).

Canebrakes (Arundinaria gigantea) too likely owed their

existence to native burning and other aboriginal land

management practices (Platt and Brantley 1997).

In the absence of aboriginal burning, massive vegetation

changes have also occurred throughout the West

(Bonnicksen 2000). Millions of acres of aspen have been lost

to invading, fire-sensitive conifers (Bartos and Campbell

1998, Kay 1997a). Similarly, vast acreages of ponderosa

pine and Douglas fir (Pseudotsuga menziesii) forests are

being replaced by more fire-sensitive, and more shade

tolerant conifers. In many areas of the West, native

grasslands have declined precipitously as fire-sensitive,

woody species have increased (e.g., Miller and Rose 1995).

Oak dominated communities in California and Oregon have

also been severely reduced by invading conifers, until today,

oak woodlands are one of the most endangered habitat

types in the West. All because native people no longer

employ fire to manage the land (Bonnicksen 2000).

Aboriginal Populations

In addition, aboriginal populations were much larger than

commonly believed (Hamel and Buckner 1998). Until

recently, it was thought that only about two million natives

inhabited North America prior to the arrival of Columbus

(Stannard 1989, 1992). Dobyns (1983), however, postulated

that native people, who were attempting to escape Spanish

exploitation in Cuba, fled to Florida in ocean-going canoes

and brought European-introduced smallpox with them to the

mainland during the early 1500s. This and other diseases, to

which aboriginal inhabitants had no immunological

resistance, then ravaged native people, reducing aboriginal

populations by >90% before the Pilgrims set-foot at

Plymouth Rock. Subsequently, Ramenofsky (1987), Smith

(1987), and Campbell (1990) tested Dobyns’ hypothesis

using the archaeological record and concluded that a major

collapse of native populations occurred ca. 1550-1600

throughout North America — 100 to 200 years prior to direct

contact of Europeans with native people in many areas; i.e.,

European diseases were transmitted from native group to

native group across all of North America — termed pandemics.

Based on this and other evidence, it is now believed that in

1492 there may have been as many as 100 million native

people in North America with perhaps an even larger number

in South America (Stannard 1992). Although Dobyns’

hypothesis is still debated (Snow 1995), in general,

estimates of pre-European native populations have steadily

been revised upward. Needless to say, prior to European

influence there were more than enough native people to

have structured vegetation communities throughout North

America (Bonnicksen 2000). This is especially true in the

eastern United States where agriculture supported high

densities of native people (DeVivo 1991, Hamel and Buckner

1998, Smith 1987).
24 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

There was no “wilderness.” There were no unnamed

streams, there were no unnamed mountains. The idea that

North America was a “wilderness” untouched by the hand of

man before 1492 is a myth, a myth created, in part, to justify

appropriation of aboriginal lands and the genocide that befell

native peoples (Cronon 1995, Denevan 1992, GomezPompa and Kaus 1992, Simms 1992). Moreover, there is no

evidence that native people ever purposefully limited their

populations to avoid environmental impacts or that, if they

tried, they were successful (Blurton-Jones 1986, 1987;

Cohen 1977, 1989; Smith and Winterhalder 1992).

Conclusions

Prior to European discovery of the New World, aboriginal

use of fire was widespread in both western and eastern

forests. In fact, the Americas, as first seen by Europeans,

had largely been crafted by native people, not created by

nature (Bonnicksen 2000, Kay 1998). Thus, the only way to

preserve original vegetation conditions in parks and other

natural areas is for modern land managers to reinstitute

historical burning regimes. A hands-off or “natural-regulation”

approach by today’s land managers will not duplicate the

ecological conditions under which eastern deciduous forests

developed. Instead, letting-nature-take-its-course creates

highly unnatural conditions that have never before existed in

eastern or western forests (Bonnicksen 2000, Hamel and

Buckner 1998). Unless the importance of aboriginal burning

is recognized, and modern management practices changed

accordingly, our ecosystems will continue to lose the

biological diversity and ecological integrity they once had

even in parks and other protected areas (Kay 1998).

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28 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Abstract

Three factors, human population density, topography, and

culture interact to create temporal and spatial differences in

the frequency of fire at the landscape level. These factors

can be quantitatively related to fire frequency. The fire model

can be used to reconstruct historic and to predict future

frequency of fire in ecosystems, as well as to identify longterm changes in anthropogenic fire regimes. Topographic

roughness is positively related by a regression equation to

the length of mean fire intervals indicating that fires are less

frequent in rough than in flat terrain during periods of low

human population density. The strength and direction of this

relationship diminishes as the frequency of anthropogenic

ignitions increases to the point that the fuel environment is

pyro-saturated. Human population density is a master

variable in understanding anthropogenic fire regimes and

topographic effects. The interactions of these factors through

time creates at least two stages in anthropogenic fire

regimes: an Ignition Limited Stage in which fire frequency is

function of human population density, and a Fuel Limited

Stage during which fire frequency is limited by fuel

production and is independent of increases in human

population density.

Introduction

Human-Landscape Interactions Affect Fire Regimes

Humans are by far the most important factors influencing the

frequency of fire ignitions in the Missouri Ozarks and

elsewhere throughout much of eastern North America. Most

fires occur during the fall, winter, and spring in years of

normal precipitation. Dry, warm weather of only a few days is

sufficient to cure fine fuels and permit the spread of surface

fires, especially in the spring and fall when dead ground

vegetation and leaf litter are most abundant and exposed to

the sun. Fires during the growing season are rare but do

occur during very hot and dry summers. High fuel moisture

content and precipitation in summer limit fire ignitions and

spread. Despite an abundance of thunderstorms in Missouri

(50-70 thunderstorm days per year) (Baldwin 1973), natural

ignitions are rare (Schroeder and Buck, 1970). Most

thunderstorms occur in the spring and summer and are

usually accompanied by heavy rainfall. Fire statistics for

Missouri (Westin 1992) indicate that less than 1 percent of

fires are caused by lightning, all other causes are humanrelated.

Topography and the fuel environment also are important

variables that determine the fire regime by controlling fire

spread, intensity and extent (Whelan 1995, DeBano and

others 1998). Human culture and land use directly affect the

continuity and nature of the fuel environment. We present a

quantitative approach to modeling the interaction of humans

with landscapes and their combined effects on fire regimes.

Three factors, human population, topographic roughness,

and culture, interact to create temporal and spatial

differences in the frequency of fire at the landscape level.

Succession in Anthropogenic Fire Regimes

Through time, fire regimes undergo a number of changes

based upon population density, topography, and culture. A

progression of four stages in anthropogenic fire regimes can

be identified in the upper Current River watershed fire record.

These include: 1) an Ignition Limited stage, 2) a Fuel Limited

stage, 3) a Fuel Fragmentation stage and, 4) a Culturally

Limited stage. During Stage 1, fire frequency increases as

the human population approaches 0.64 humans/km2

 and the

number of potential ignitions increases. Fire frequency is a

logarithm function of human population density. In later

stages, as human population density increases and the

landscape is saturated with ignitions, other factors limit fire

frequency such as fuel loading and continuity, cultural

attitudes toward fire, and human valuation of natural

resources. In the Current River watershed, the second stage,

a Fuel Limited Stage, is characterized by limits imposed on

the frequency of fire frequency by the primary production of

fuels. During this stage, human population density exceeds a

threshold of 0.64 humans/km2 and increases in population

density and ignitions do not result in increased fire frequency.

This paper will integrate population density and topographic

roughness in an empirically derived model that can be used

to reconstruct past or to predict future frequency of fire in

ecosystems at specific sites. Here, we focus on the

development of an empirically based model of fire frequency

from over 2,500 tree-ring dated fire scars at 29 sites in the

Current River watershed in southeastern Missouri. This data

set is used to develop and verify a regression equation that

predicts fire frequency based on topographic and population

variables. We also examine the interactions of humans and

landscapes with respect to fire regimes, and illustrate, using

the fire model, the spatial and temporal variability in the

anthropogenic fire regime in the upper Current River

landscape.

Rational and Use for Fire Regime Models

Quantitative models of fire regimes can be used in many

ways. Past fire regimes can be reconstructed for natural

areas that have no on-site fire history information. The

equations can be used in conjunction with soil, geology, and

species data to reconstruct past and potential flora and

Humans, Topography, and Wildland Fire: The Ingredients for Long-term

Patterns in Ecosystems

Richard P. Guyette1 and Daniel C. Dey2

1Research Associate Professor, The School of Natural

Resources, 203 ABNR Building, University of Missouri,

Columbia MO 65211. email: guyetter@missouri.edu

2Research Forester, USDA Forest Service, 202 ABNR Bldg.

Columbia MO 65211. email: ddey@fs.fed.us
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 29

faunal mosaics. Thus, land managers interested in returning

small isolated parcels of land, natural areas, or whole

ecosystems to pre-European conditions using prescribed fire

can use this model to reconstruct the fire regime.

Researchers can also use the equation to create a GIS

overlay of a fire regime for use with research plots or to

make inferences about future fire regimes based on changes

in population and climate, and landscape variables. Perhaps

the most important aspect of fire regime models and fire

history however, may not be the information provided on fire

regimes, but the perspective provided on the long-term

interactions between humans and their environment.

The Historical Context of Human Populations,

Cultures, and Fire in the Current River Watershed

All of the historic changes in human populations due to war,

migration, and disease are probably reflected in the fire scar

record. Europeans introduced diseases decimated Native

American population. War among peoples caused migration

and genocide. Depopulation caused by disease (Dobyns

1983; Ramenofsky 1987), warfare, and migration occurred

throughout much of North America (DeVivo 1991) and has

been linked with abrupt changes in fire history (Guyette and

Dey 1995a; Dey and Guyette in press). Indeed, often the

highly variable nature of human populations and culture is

reflected in the extreme variability of the fire record within a

site (Guyette and Dey 1995b, Cutter and Guyette 1994,

Guyette and McGinnes 1982).

Anthropogenic ignitions were probably frequent along the

rich fertile bottom lands of large rivers including the Current

and Jacks Fork Rivers. About 1000 A.D. (Lynott 1989, Price

and others 1983) during the Mississippian Cultural Phase,

agricultural communities were located in riparian areas in the

Current River watershed, and they would have been a

source of many ignitions. With the end of this phase (about

1350 A.D.) began an era of limited use of the watershed by

humans from downstream reaches of the river system (Price

and others 1976). An early source of anthropogenic ignitions

may have been the 6,000 Quapaw (Baird 1980) that lived to

the southeast of the Current River in Arkansas near the

confluence of the White, Arkansas, and Mississippi Rivers

before 1680. Disease came first to large riverine cultures

such as the Quapaw because of their location and

population density (Dobyns 1983), and reduced the Quapaw

population by two thirds in 1698 and again reduced their

population in 1747 and 1751 (Baird 1980). There were only

about 700 Quapaw by 1763. Coincident with a low

percentage of sites burned circa 1750 in the Current River

watershed (Guyette and Cutter 1997) were epidemic disease

in 1747 and 1751 which reduced the Quapaw population.

The Osage people frequented the study site about the time

of first European contact and probably provided many

ignitions during the 1700s (Guyette and Cutter 1991). Banks

(1978) commented that there were about 6,000 Osage circa

1800 in and around the Ozarks. About 1,200 Great Osage

lived on the Osage River 280 km west of the Current River

(Marriott 1974). A change in the territorial boundaries of the

Osage from the late 1600s to about 1803 (Bailey 1973,

Wiegers 1985) is coincident with long-term trends in fire

frequency in the Current River watershed (Guyette and

Cutter 1997). Territorial expansion by the Osage from 1673

to 1770 probably resulted from many factors, one of which

included the acquisition of equestrian technology from

aboriginal trade with western tribes. The horse gave the

Osage new range and mobility to hunt, exploit, and culture

(by fire) areas such as the Current River watershed which

were distant from their territorial focus. Wiegers (1985)

estimated that the Osage acquired horses as early as 1680

while Waldman (1985) sets the date circa 1719. They hunted

in the east for bears (perhaps in the Current River

watershed) and in the west for buffalo (Chapman and

Chapman 1972). Stevens (1991) reported that the Osage

moved south and east toward the Current River on extended

spring hunts coincident with Missouri’s largest fire season

(Westin 1992).

The Cherokee were one of several tribes that migrated into

the Ozarks after being displaced from their eastern lands by

Euro-Americans (Gilbert 1996). They probably continued the

traditional burning practices they had used on their ancestral

lands. About 6,000 Cherokee (Gilbert 1996) may have been

living in southeast Missouri and northeast Arkansas at the

time (1803) of the Louisiana Purchase. In the late 1770s,

many of the Cherokee settled to hunt and farm along the St.

Francis River, which lies about 60 km east of the Current

River. The Osage made war upon the Cherokee (Banks

1978) as the Cherokee infringed upon their hunting grounds.

Conflict may have led to wildland burning. A general increase

in the percentage of sites burned annually (Guyette and

Cutter 1997) in the Current River watershed occurred from

1760 (9 percent) to 1820 (30 percent) coincidently with the

migration of the Cherokee (Gilbert 1996) and other eastern

tribes.

The Delaware had a tradition of wildland burning before

migrating to Missouri from Ohio (Whitney 1994). The

Delaware and Shawnee entered Missouri at Cape Girardeau

and passed through the Current River watershed on their

way west in the late 1700s and early 1800s (Stevens 1991).

One estimate of Shawnee and Delaware population west of

the Mississippi in 1812 was 400 (Marriott 1974). In 1816,

Banks (1978) stated that there were about 840 Delaware

and 1,300 Shawnee in all of Missouri. In 1824 there were

1,383 Shawnee in Missouri (Howard 1981). Delaware

hunted, lived, and traded in the upper Current River area

from about 1815 to 1822. In November 1820, 1,346

Delaware crossed the Mississippi River and made an

emergency encampment on the Jacks Fork of the Current

River (Weslager 1978). However, they stayed a number of

years and this was associated with increased fire frequency

(Guyette and Dey 1997). Most of the Delaware and

Shawnee had left the Current River watershed by the 1830s

and Euro-Americans began to occupy the area.

Old-stock Euro-Americans (Gerlach 1986) from the

southeastern U.S., mainly Tennessee, began settling in the

area circa 1820 and continued their tradition of wildland

burning. Later, circa 1860, Scotch-Irish immigrated into the

area. The Current River watershed, like many other forested
30 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

areas in the eastern United States (Sutherland 1997),

experienced an increase in fire frequency with settlement by

Euro-Americans. In the Current River watershed the percent

of sites burned increased with population density during the

1810 to 1850 period and the average percent of sites burned

in a year nearly doubled from 20 percent in 1810 to 39

percent in 1850 (Guyette and Cutter 1997).

Methods

Fire Scars

Fire scars on survivor trees form the data base of this work.

In anthropogenic fire regimes, fire scars can be thought of as

ecological artifacts. They are objects made or modified by

humans. Humans set wildland fires and wounded these

woody plants as they attempted to manipulate vegetation

with fire for a variety of purposes. As ecological artifacts,

they point out the relationships between humans, organisms,

and forests. Like artifacts in an archeological context, they

can be dated in time and given a location in space. As such,

they reflect many human characteristics such as population

density and culture that can be used in an anthropological

context. The dates and locations of these ecological artifacts

are used in this work to examine the relationships between

humans, fire, and landscape over a period of extraordinary

change and diversity in human history.

The regression equation for reconstructing historic fire

frequency is derived from data on fire frequency, topographic

roughness, and human population density. The mean fire

intervals (MFI) used in the regression equation were derived

from fire scar data for three time periods (1700 to 1780, 1781

to 1820,1821 to 1850) at the 29 sites. Much of the fire history

information that follows is compiled from more than 2,500 fire

scars on shortleaf pine (Pinus echinata Mill.) stumps, trees,

and remnants in the Current River watershed (Guyette and

Cutter 1997, Batek and others 1999). Dendrochronological

methods were used to date the fire scars found on over 166

shortleaf pine which are used as the basis for the model and

following discussion. The mean fire intervals (MFI) in this

study are thought to reflect fire frequency during ecosystem

development and a range of human population densities that

has occurred over at least the last millennium in the Ozarks.

Here, we present the two most significant variables in an

equation that characterizes the effects of human population

density and topographic roughness on the frequency of

wildland fire.

Human Population Estimates

A spatial and a temporal component of human population are

combined to reconstruct human population density values for

use in the equation. The spatial component reflects the

tendency in the watershed for greater population densities

and therefore ignitions to occur downstream. We use the

distance upstream of a site from Hawes, on the Current

River, as a reference variable we call River Mile to model this

tendency. Many archeological and historic Native American

sites are located in the larger and more fertile bottom lands

in the downstream reaches of the Current River (Stevens

1991). For instance, a major Osage trade route to St. Louis

intersected the lower Current River. The abundance of these

sites decreases significantly (r=-0.53, p=0.03) with their

distance upstream. The Mississippian cultural phase

(O’Brien and Wood 1998) persisted downstream on the

southeast border of the Ozarks until as late as 1700 (Price et

al. 1976). Quapaw lived downstream and southeast of the

watershed (Rafferty 1981). Later, the density of human built

structures in the Current River watershed during the mid

1800s diminishes with their distance upstream (r=-0.69,

p=0.002). Even today the population of towns along the

Current River and Jacks Fork Rivers decreases as one

moves upstream (r=-0.98, p=0.02). Perhaps fertile soils of

the lower reaches of the river produces more fuel and

forage, which in turn attract more game, hunters, gatherers,

and agriculturalist. These are relationships that may persist,

to some extent, through time and cultures. Alternatively,

fertile soils may be correlated with landscape level fuel and

fire dynamics, or land form gradients such as topographic

roughness, that may affect the ignition, spread and

consequently the frequency of fire.

We derived the temporal component of human population

density in the study area from population numbers and

trends as described in historical accounts and analyses as

reviewed in the introduction. Fire frequency has been found

to be strongly associated with human population density

(Guyette and Dey 1997, Dey and Guyette in press, Guyette

and Cutter 1997). The density of each group in the Current

River watershed was calculated by dividing their total

population by the area of a circle whose radius was the

distance between the population center and the study site.

This component of human population density does not take

into account the great spatial variation of population density

within an area and only reflects changes in population

density over time at study sites as a whole.

An Index of Topographic Roughness

The highly dissected topography of the Current River

watershed has been shown (Guyette 1995) to affect the

spread and frequency of fires during periods of low human

population density when anthropogenic ignitions were limited

in number. Roughness can slow a low intensity fire by

decreasing the rate of spread as fires burn down steep

slopes, fuel continuity is broken by creeks, rivers and rocky

outcrops, and changes in aspect influence fuel moisture.

Topographic roughness is estimated by measuring the

surface area of the earth with measuring surfaces of different

sizes (Figure 1). Indices of topographic roughness are

developed by comparing surface area measurements made

with two different sized scales. A circle, 5,000 meters in

diameter, is marked on a digital elevation map. The surface

area of the earth enclosed by this circle is calculated from

pixels that are 30 meters on a side. Their slope and a

trigonometric conversion are used to estimate the area of the

uneven land surface (Krstansky and Nigh 1999). The pixels

are summed to estimate the surface area of the landscape

enclosed by the circle. This measure is then divided by the

planimetric surface area (the large scale in this case) of the

circle that is 5,000 meters in diameter. This ratio of the actual
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 31

surface area to the planimetric surface area is the Index of

Topographic Roughness.

Results and Discussion

Modeling Fire Frequency

We use the Index of Topographic Roughness and estimates

of human population density to model the frequency of fire at

the landscape level. Central to this model are regression

equations derived from and tested with independent fire scar

data. The regression used to reconstruct past fire frequency

at the study sites for the period 1700 to 1850 is:

MFI = -442.1 + (449.9 x topo) -(0.001 x pop), (Equation 1)

where: MFI= mean fire interval

topo = Index of Topographic Roughness for a circle

with a 5,000 meter diameter

pop = natural log of human population density times

the square of river mile,

n = 78 (three time periods, 29 sites),

r

2

 = 0.51,

all variables and intercept are significant at P<0.01.

This regression explains about half of the variability in fire

frequency in time and space. The amount of unexplained

variability is not surprising considering the large role chance

plays in a complex and highly variable phenomena like

wildland fire. In addition, there are methodological problems

with the fire scar record such as the number of sample trees

at a site, their location, the size of the site, and the hit and

miss nature of the scarring process that create errors with

respect to the actual spatial and temporal distribution of fire

events in the data set. None-the-less, the temporal and

spatial depth of the fire scar record allow for the identification

of the significant independent variables in this equation.

Although the framework of this equation may be applicable in

many ecosystems, the calibration of these variables will

differ greatly in different ecosystems. Differences in fuels,

climate, topographic roughness, and other landscape

variables such as the abundance of lakes would require a

new calibration.

Human Population Density and Topographic

Roughness in Fire Regimes

Anthropogenic fire regimes are the result of complex

interactions between human population density and an

number of factors, such as culture, fuels, and landscape.

Topographic roughness is one of the most important and

temporally persistent landscape variables. At low population

densities, fire frequency increases as population density

does as a result of increasing anthropogenic ignitions (Figure

Figure 1.—Topographic roughness is an important variable which interacts with human

population density to control the frequency of fire. Topographic roughness can be estimated

with an Index of Topographic Roughness. The illustrated example of an Index of Topographic

Roughness is calculated by measuring the circumscribed region of the earth of a given

dimension and shape with a large measuring surface ( ls ) and many small measuring surfaces

(ss

). The sum of the areas (Sa) of the small measuring surfaces is then divided by the area (La)

of the large measuring surface to obtain the Index of Topographic Roughness.
32 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

2) and topographic roughness is an important factor

mitigating the frequency of fire (Figure 3). At higher

population densities fire frequency may be fuel

dependent or diminish as landscape artifacts (i.e., fuel

continuity or mosaic that result from human land use),

culture, and changes in human economics (ex. real

estate, forage, timber) inhibit the ignition and

propagation of fire over the landscape. The effects of

topographic roughness on fire frequency change with

time and human population (Figure 4). Topographic

controls on the frequency of fire become less and less

important as population density and the frequency of

anthropogenic ignitions increases. Just before the era

of fire suppression (Figure 4), as population density

and agricultural activity become greater, burning

increases in topographically rough areas relative to

topographically smooth areas as humans attempt to

culture vegetation in areas of marginal agricultural

productivity. A rule-of-thumb concerning fire history is

that at low population densities topographically smooth

areas (often prairies and grass lands) burned more

often than topographically rough areas (often forested

lands). As population density increases,

topographically smooth areas burn less often and

topographically rough areas burn more often.

Pyro-saturation is a stage in a fire regime when fuels

are burned as soon as they accumulate enough to

carry a fire. Inorganic decomposition (fire) plays a

much greater role than organic decomposition (decay)

in controlling the oxidation of biomass. Ecologically,

the resistence of an ecosystem to pyro-saturation is a

function of human population density and topographic

roughness. The resistence of some ecosystems to

pyro-saturation from anthropogenic ignitions is very

low. For instance, it might take only a handful of

humans on horseback or foot to keep a topographically

smooth ecosystem, such as a large prairie of tens of

thousands of square kilometers, in a state of pyrosaturation. On the other hand, many topographically

rough ecosystems, such as the forests of the Central

Hardwoods Region, require a relatively high human

population density to reach and maintain pyrosaturation. The quantitative relationships expressed in

Equation 1 allow us to estimate that in the Current

River watershed it takes about 0.64 humans per

square kilometer to reach a state of pyro-saturation.

Implications for Landscape Fire Regime

Diversity

The fire model (Equation 1) was used to predict the

mean fire interval for points throughout the upper

Current River watershed (Figure 5). The value to this

approach to studying landscape-level fire regimes is

that extrapolation of single fire histories of limited

spatial and temporal dimensions often underestimate

the actual variability within an ecosystem. In contrast,

a landscape fire model can represent the diversity of

fire regimes as they change across the landscape with

variations in topographic roughness and human

Figure 2.—Human population density is the major factor in the

frequency of wildland fire ignitions in many regions. This scatter

plot and regression illustrate the change in the percentage of

study sites burned each year with human population density.

The data is for the period 1620 to 1850 and is from the Current

River watershed in Missouri. The percentage of sites burned (y

axis) is from annual data that has been smoothed with an 11

year moving average (Guyette and Cutter 1997).

Figure 3.—These two scatter plots and regression lines illustrate

the mitigation of fire frequency (MFI) by topographic roughness

during two periods of low human population density.

Topographic roughness is hypothesized to have reduced the

frequency of fire by inhibiting the ignition and spread of

anthropogenic fires.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 33

population. The diversity of mean fire intervals present

at the landscape level between 1700 and 1780, a

period of low population density, may best reflect the

fire regime as it was for many centuries before

European settlement.

The diversity of fire frequency at specific locations in

the Current River watershed is much greater at low

population densities than at high population densities

as the spread of fires from a limited number of

anthropogenic ignitions is mitigated by topographic

roughness. The spatial diversity of fire frequency within

an ecosystem that results from topographic roughness

persists through time and can result in strong species

associations. Long-term topographic control of fire

regimes can create refugia for fire sensitive species in

topographically rough areas as well as creating refugia

in topographically smooth areas for fire dependent

species. These refugia may provide valuable sources

for re-colonization of disturbed landscapes.

Conclusions

Fire frequency varies spatially and temporally at the

landscape-level largely due to changes in topography,

human population, and culture. Population density and

topographic roughness are master variables in

understanding anthropogenic fire regimes that can be

quantified and used to model the variability within a fire

regime. Although landscape level fire models require a

considerable investment in the acquisition of fire scar, site,

landscape, and population data, they enable an

understanding of long-term fire regimes that can be used in

many ways.

Our fire model can be used to estimate the frequency of fire

at any or all points in a landscape. Fire frequencies and

regimes can be calculated for natural areas, parks, and

forests that have no fire scar, historic, or charcoal data base

for estimating a fire regime. Future predictions of fire

frequency can be made by incorporating changing human

population density, values, climate, and fuels into the model.

The model enables the mapping of variation within a fire

regime in both time and space. Maps of fire regimes and

estimates of the long-term frequency of fire can be used in

research on flora and fauna, in setting the frequency of fire in

prescriptions to restore or maintain fire-dependent

ecosystems, and the development of silvicultural

prescriptions that mimic “natural” disturbances. Maps of fire

regimes and the long-term frequency of fire disturbances can

also be used to identify areas in the landscape that are

refugia for either fire-dependent or fire-sensitive species.

Anthropogenic fire regimes, composed of complex

interactions among fire, humans, fuels, and topography,

have altered a continuum of ecosystems in North America for

many thousands of years. These fire regimes are dynamic

and have a number of identifiable successional stages that

change as humans populate and transform their environment

by fire and artifact. Human-induced changes in a fire regime

are modified by topography and fuels. Stages in

anthropogenic fire regimes are marked by changes in factors

which limit the frequency of fire such as the frequency of

ignitions, the production of fuels, the propagation of fire, and

changing cultural values.

Acknowledgments

The authors thank the Missouri Department of Conservation,

the Missouri Ozark Forest Ecosystem Project, the National

Park Service, and the USDA Forest Service for their support

of this research effort.

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Figure 4.—This graph illustrates how human population density can

mitigate the effects of topographic roughness on the frequency of fire.

The y axis is the correlation coefficient between fire frequency (MFI)

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34 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

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Figure 5.—This map illustrates the diversity of

mean fire intervals present at the landscape level

during a period (1700 to 1780) of low to

moderate population density, which best reflects

the fire regime many centuries before European

settlement. Fire frequency gradients are

represented by the iso-pyro lines. Iso-pyro lines

define a gradient of equal fire frequencies.

Numbers adjacent to iso-pyro lines give the class

of mean fire intervals defined by each line.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 35

Guyette, R.P. 1995. A tree-ring fire history of wildland fire

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Ph.D. Dissertation. University of Missouri, Columbia, MO.
36 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Abstract

Unlike many areas of the United States, anthropogenic fires

are the prime agent for affecting changes in plant and animal

species composition in the southern Appalachian Highlands.

Although the extensive use of fire by the American Indians

has been recognized from the earliest European observers,

it is somewhat difficult to determine the impact prehistoric

fires had on forest structure. By examining the fossil and

charcoal record from Cliff Palace Pond with the

archaeological record recovered from nearby prehistoric

sites, a 9,500-year record of the vegetational development

can be established for the escarpment zone of Eastern

Kentucky. This record indicates that anthropogenic fires

played a central role in shaping the forest structure,

especially after the transition from a hunting and gathering

economy to one based on swidden agricultural practices.

Introduction

Most federal land managing agencies are now committed to

using ecosystem-based approaches in their land

management objectives. Unfortunately, until recently, these

approaches were primarily concerned with the physical and

the biological components of the ecosystem and usually

lacked a comprehensive understanding of the human

dimension within the environment. Yet the management of

federal lands is not set by nature, but by people (Williams

1993). Just as people shape the management direction of

ecosystems today so has it been since the first Paleoindians

set foot on the New World. Here in the eastern woodlands,

people have been a part of ecosystems since the end of the

Pleistocene and have been shaped by, and in turn shaped

ecosystems in a number of ways.

On the threshold of the 21st century we are striving as never

before to bring a historical context into natural resource

management. At long last both land managers and the public

are beginning to accept that there is a historical relationship

between people and nature, that all human history has a

natural context and that we shape the land just as much as

the land shapes us (MacCleery 1998). Wildland fires (both

natural and those created by people) have long been

considered to be an integral part of ecosystems across North

America and that the recurring fire disturbances are essential

to the functioning of these systems (Mutch 1995). While fire

certainly played a role in shaping the forests of the southern

Appalachians, the extent that anthropogenic fires played has

been poorly understood.

Cumberland Plateau Escarpment Zone

In Kentucky, conglomeratic sandstones outcrop in a belt

ranging from 8 to 40 km wide along the western edge of the

Cumberland Plateau. Geologically, this area is referred to as

the Pottsville Escarpment. Erosion within this belt has

created deep narrow stream valleys flanked by steep-sided

to precipitous valley walls. Little level land is available along

the stream margins but the ridges, ringed by high sandstone

cliffs are fairly wide and flat. Nearly 5000 kilometers of

cliffline snake through the Daniel Boone National Forest, and

often stretch for miles without a break. These natural

features led Miller, in his Geology of Kentucky (1919) to refer

to the escarpment zone as a “Chinese Wall” that greatly

inhibited economic development of the area.

According to a tree census of eight “old forest” stands in the

Lower Big Sandy River drainage, the dominant trees of

Eastern Kentucky were several oak species (especially the

white oak), beech, hickories, pines, maple and poplar

(Shaler and Crandall 1876:12-13). Along the narrow stream

valleys cane thickets was the dominant understory. Nuttall

(1821:29) noted that the northern limit of cane was the

southern bank of the Ohio River. It is from the archaeological

sites and the natural ponds within this narrow escarpment

zone that an understanding of prehistoric fire and its affects

on forest structure are beginning to emerge.

Fire History

Wildland fire in the southern Appalachians results from two

forces. One is natural, those caused by lightening and the

other cultural, those which result from people. Martin

(1990:56) reports that along the Cumberland Escarpment,

lightening-caused fires account for only two percent of the

total fires documented within a 50-year period. Here, the

annual lightning fire occurrence is less than five fires per one

million acres and usually occur after the month of May when

the sap is up in woody plants. Consequentially, these fires

usually lack the intensity necessary to affect plant species

composition.

Since the natural fire regime was not sufficient to account for

the fire-adapted ecosystems noted by early observers it was

the anthropogenic use of fire that shaped the ecosystems of

the eastern woodlands more so than any other force. Fire

ecology has been practiced to various degrees for the last

hundred years within the pine forests of the southeast.

However, resource management agencies have been

reluctant to use anthropogenic fires (prescribed fire) as a tool

for managing the central hardwood forests. It has only been

within the last decade that prescribed fire in hardwood

forests has become accepted and this acceptance is far from

universal.

Fire on the Edge: Prehistoric Fire Along

the Escarpment Zone of the Cumberland Plateau

Cecil R. Ison1

1Forest Archaeologist, USDA Forest Service, Daniel Boone

National Forest, 1700 Bypass Road, Winchester, KY 40391
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 37

A baseline in the fire history of the Cumberland Plateau must

begin with the historical literature. Although the

pervasiveness of anthropogenic fire makes it a nearuniversal instrument for analyzing the interaction of people

with their environments (Pyne 1994) without the written

record as a guide, many of the material conditions observed

in the archaeological record may not become apparent.

Therefore we must rely on the observations of early writers

to provide the foundation of aboriginal fire practices.

Wherever early explorers journeyed they remarked on the

Indians use fire. These ranged from burning the woods to

clear land for cultivation to providing herbage and browse for

the white-tailed deer. The deer were considered the cows of

the eastern woodland Indians and the burnt woods their

pastures (Whitney1994: 109). It has been estimated that this

species alone supplied up to 90 percent of the animal protein

consumed by the prehistoric inhabitants of eastern Kentucky

(Wyss and Wyss 1977:24).

Merely 36 years had lapsed from Columbus’ first sighting of

the New World until Cabeza de Vaca recorded the first

description of the Indian methods of burning the woods

(Favata and Fernandez 1993:72). From de Vaca’s first

written account onward through the 19th century, the historic

literature is full of descriptions of Indians burning the woods.

Travelers repeatedly remarked on the wildfires they

witnessed during the initial settlement of the Ohio Valley in

the mid 1700’s to early 1800’s. In “Images of the Ohio Valley”

historian John Jakle (1977) relates how early travelers

marveled at the openness of the forest floor and its almost

park-like appearance within the oak, hickory and chestnut

communities. Fires according to Jakle, “provided the most

spectacular forest scenes. Fires burned out of control each

fall charring hundreds of thousands of square miles...over a

distance of fifty miles the flames reflected on the clouds of

smoke (1977:54).”

The earliest historical accounts of wildfire along the

escarpment of the Cumberland Plateau were those of Dr.

Thomas Walker, an explorer for the Loyal Land Company of

Virginia. In 1750, Walker set out from Charlottesville, Virginia

to inspect the western country to select the location of the

company’s 800,000 acre land grant. The first reference to

forest fires by Walker was on May 17, 1750 in the vicinity of

the divide between the waters of the Rockcastle and South

Fork of the Kentucky River in what is now Jackson County,

Kentucky. On May 17th he penned in his journal “The woods

have been burnt some years past and are now very thick,

the timber being almost kill’d” (Johnston 1898). Ironically, this

is in the same vicinity of Cliff Palace Pond. The only other

reference to wildfire that Walker made during his journey was

again along the escarpment. On May 30th while traveling

along the headwaters of the north Fork of the Red River

Walker notes “The woods are burnt fresh about here and are

the only fresh burnt woods we have seen these six weeks”

(Johnston 1898).

Sometimes the clues of wildfires come from the most tragic

accounts of the early Kentucky pioneers. The rescue party

searching for the victims of the 1793 Easter Sunday attack

on Morgan’s Station for example, noted that two of the

victims had been forced to march through a recently burned

section of the woods. James Wade recounts that on April 2nd

“...just above the head of Little Slate we found Mrs. Becraft

and her suckling child ‘six or eight months old’ lying

tomahawked. It was a very plain case. They had marched

her that far in her shift, as was visible from the scratches and

marks on it from a burnt wood they had passed through. And

there she had given out” (Hogan 1991:24-25). This location

is very near a large rockshelter known as Newt Cash Shelter,

which would later provide the first archaeological evidence of

prehistoric anthropogenic use of fire.

With the hundreds of accounts penned by early observers

we have no problem accepting the purposeful use of fire by

American Indians for a multitude of purposes during the

historic era. In fact, it is generally agreed that Native

Americans had a sophisticated knowledge of the use of fire

(Patterson and Sassaman 1988). The problem arises when

we try to provide time depth to the use of fire before the

written record. After all, time depth is essential in the overall

understanding of the role of fire in ecosystems. Despite this

documentary evidence many still hold the view of the early

pristine forest, one untouched by the hand of man, where

native people lived in the forests without changing the

ecosystem in any way. Where do we find the proof to

counteract this view? It comes from unlikely sources. It

comes from fire-scorched ancient trees, it comes from

natural ponds whose sediments hold the record of nearby

plant life from the first colonizers at the bottom to the current

tenants at the top and it even comes from people’s bowel

movements preserved in the dry environments of caves and

rockshelters over the millennia. But above all else, the proof

comes from an interdisciplinary approach. It comes from the

cooperative efforts of natural and heritage resource studies.

Archaeological Evidence

Archaeological investigations are essential for providing a

diachronic context into the anthropogenic role of fire. In

Kentucky, when the field of modern archaeological research

was just beginning to become established in the late 1920’s

(Lewis 1996) early archaeologists realized the importance of

the dry rockshelters and caves in recording the story of the

human saga. Along the escarpment archaeologists

recovered textiles, grass beds, desiccated human feces and

other perishable artifacts from the dry rockshelter deposits.

These materials provided a wealth of detail to our

understanding of the lives of Kentucky’s early inhabitants

and their effect on landscape ecology. The following brief

descriptions of the investigations at four of these

rockshelters (figure 1) provide a tantalizing glimpse into

anthropogenic fire behavior. The recent analysis of the Cliff

Palace Pond sediments has provided a further dimension on

fire history studies within the central hardwood region.

Newt Kash Shelter

The earliest clues for the prehistoric use of anthropogenic

wildland fire in Kentucky began in the 1930’s when Volney

Jones, an ethno-botanist at the University of Michigan

examined a small sample of vegetal remains recovered the
38 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Newt Kash rockshelter site in Menifee County. Most of the

remains analyzed by Jones were from the sleeping beds of

the aboriginal inhabitants. These beds were made of a matlike fabric covering lain over a bed of grass, leaves and straw

(Webb and Funkhouser 1939). The bedding also contained

all sorts of miscellaneous paraphernalia, which had been

accidentally incorporated into the grass mat. Among these

were well-preserved seeds, nuts, textile refuse, wood,

charcoal and fecal material.

Jones’ analysis indicated that the grass, which made up the

bulk of the bedding was big bluestem (Andropogon furcatus),

a robust grass commonly associated with prairie or open

woodland environments. Of the identified woods, hickory

was the most common followed by chestnut and oak, again

species that are uniquely fire adapted. Other trees and

shrubs included ash, box elder, sumac and poplar (Jones

1936:160). What intrigued Jones the most about this sample

was the preponderance of cultigen seeds including

goosefoot (Chenopodium sp.), sunflower (Helianthus annus),

squash (Cucurbita pepo var. melopepo), marsh elder (Iva

sp.) and canary grass (Phalaris caroliniania).

The paleofecal samples analyzed by Jones also were also

quite intriguing. All the cultigens identified within the general

contexts of the site with the exception of squash, were

represented with the fecal matter. Sunflower for example,

while found sparingly elsewhere in the materials examined,

comprised a high percentage of the fecal bulk. When

combined with the goosefoot, marsh elder and canary grass

seeds, plants which were foods grown by simple agricultural

methods, comprised the majority of the bulk of the fecal

matter.

The study of the ethno-botanical materials from the Newt

Kash Hollow rockshelter led Jones to speculate the

occupation of the shelter was during a period of transition of

the flora from prairie to forest or reverse (ibid p.165).

Although archaeologists in the 1930’s had no way of

absolutely dating the cultural deposits, Jones felt tempted to

place the occupation of the site around 4000 years ago. With

the advent of radiocarbon dating in the 1950’s the intensive

occupation of Newt Kash shelter has been determined to be

circa 3,000 years ago (Smith and Cowan 1987, Gremillion

1997). Never the less, Jones’ work was a keystone for the

use of botanical remains from archaeological sites in

examining floristic changes upon the landscape.

Cloudsplitter Shelter

It would be over forty years before the well-preserved

materials beneath the secluded cliffs of eastern Kentucky

once again gained prominence. Although the information

gleaned by the early excavations of the 1920s and 1930s

had been used to formulate hypotheses concerning the

development of horticulture in eastern North America, the

precision of the early excavations made it difficult to address

synchronic research topics (Cowan and others 1981:60).

This changed in 1978 when the National Science Foundation

funded excavations at the Cloudsplitter Rockshelter. One of

the specific goals of the excavations was to address how

human cultural disturbance, especially early horticultural

practices affected or altered the local environment. This

protected overhang, perched 250 meters above the deeply

entrenched Red River floodplain in Menifee County,

Kentucky provided the first well-recognized quantitative data

sets on early horticultural transitions within the eastern

woodlands

The careful excavation of the complex cultural layers at

Cloudsplitter provided a 9000-year record of human

habitation and interaction with their local environments. At

the time of initial occupation the shelter circa 7,000 B.C. the

people were faced with a still-evolving postglacial landscape

where a dense stand of hemlock shielded the front of the

overhang. Today, the mature hemlock first occurs midway

down the slope from the site. However, by 1,000 B.C. the

deposits of Cloudsplitter, like Newt Kash began to reveal

anthropogenic modifications of the local floristic

communities. Non-economic plant remains recovered from

the Late Archaic deposits (circa 1,000 B.C.) which are

indicative of a fringe environment in an otherwise closed

Figure 1.—Escarpment Zone sites discussed in the text.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 39

canopy forest led Cowan (1985:330-343) to speculate

that areas within the immediate vicinity of the shelter

had been cleared away. The several cultivars

recovered from the Late Archaic deposits including

squash (Cucurbita pepo), gourd (Lagenaria siceria),

sunflower (Helianthus annuus), and goosefoot

(Chenopodium berlandieri) probably reflects upland

garden plots within the clearings (Ison 1991).

Cold Oak Shelter

In 1984, the Forest Service undertook salvage

excavations at the severely vandalized Cold Oak

Shelter located at the head of a remote hollow in Lee

County, Kentucky. While heavily disturbed by looters,

the investigations documented two cultural

occupation zones containing a suite of cultigens

similar to those recovered from Cloudsplitter and

Newt Kash. The pattern that began to emerge from

Newt Kash and Cloudsplitter was repeated at Cold

Oak. Once again, the earliest component containing

the cultivars was a Terminal Archaic horizon dated to

circa 1,000 B.C. (Ison 1988, O’Steen and others

1991). It was beginning to become apparent that by

the first millennium B.C. the forests surrounding these sites

were undergoing major changes at the hands of people.

Ten years later, under the sponsorship of the National

Geographic Society, more in-depth excavations were

undertaken at Cold Oak (Gremillion 1995, 1998). Building on

the earlier investigations, the focus of 1994 data recovery

project was to examine the changes in the ecological

relationships between people and the plants under their

management (Gremillion 1998). The investigations revealed

that the quantities of seeds of cultivated plants increased

dramatically in the post-1000 B.C. deposits concomitantly

with the decrease of seeds represented by taxa of forested

habitats (Gremillion 1998). In other word, the analysis of the

seed data indicated a general increase in anthropogenic

habitats such as gardens and clearings near the habitation

sites (figure 2).

Courthouse Rock

The pattern of early plant domestication along the

escarpment of the Cumberland Plateau was once again

substantiated with the investigations at Courthouse Rock, a

sandstone overhang situated approximately 5 kilometers

west of Cloudsplitter. The normal suite of seeds representing

cultivated plants were recovered from cultural deposits

radiocarbon dated between circa 500 and 1000 B.C. While

specimens of cultivated plants made up the largest

percentage of the identified seeds, the overall assemblage

indicated a canopy opening near the site that permitted

colonization by sun-loving plants (Gremillion 1999:48). This

is most likely the result of preparation of upland garden plots.

Green Sulphur Spring

Even open sites that do not maintain the preservation

characteristics of dry rockshelter sites can provide data on

anthropogenic forest changes. At the Green Sulphur Spring

Site Complex in southern West Virginia differential wood

types were noted between the Middle Woodland component

(circa A.D. 200 – 400) and the Late Woodland Component

(circa A.D. 400 – 800). In sharp contrast to the dominance of

hardwood charcoal (99.3 percent) within the Middle

Woodland deposits pines represent over 83 percent of the

Late Woodland charcoal. This shift in forest composition is

thought to have been the result human-induced changes due

to land clearing associated with horticulture. As the fields

became exhausted or overgrown after several years of

cultivation or from periodic abandonment of the site, the

fields reverted to forest again with pine being the initial

succession species (Rossen 1985, Rossen and Ison 1986).

Hill-side Farming

If these sites are accepted as providing a reasonably

accurate indication of prehistoric horticultural endeavors,

then the next question is to what extent fire played in the

clearing of areas for forest farming. Although there are

numerous conditions affecting the decision making process

that went into the placement of Kentucky’s first garden plots,

there are two major underlying cultural mechanisms

responsible for their location. These are the “Slug Principal”

and “Jones’ Law”. The Slug Principal dictates that there will

be a gravitation toward least effort to produce a food crop

(Binford 1983:200-201) while Jones’ Law states that “with

any given society, the amount of energy obtained in overall

subsistence activities must equal or exceed the amount of

energy expended in those activities’ (Cowan 1988:459).

Given these two assumptions, combined with the technology

available at the time of early plant domestication, the

conditions that most greatly affected development of early

horticultural practices along the Cumberland Escarpment

were the local topography and the use of fire.

Figure 2.—Forest and disturbed seed taxa from Cold Oak

shelter cultural deposits. Adapted from Gremillion 1998.
40 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Along the escarpment, little level land is available along the

stream margins and where available it is covered by a

luxuriant growth of moisture tolerant species. These included

the dense stands of cane (Arundinaria sp) mentioned by the

early explorers of Eastern Kentucky (Filson 1969:318).

During the Terminal Archaic Cultural Period (circa 1500 –

500 B.C.) the digging stick was the tool of use, not the hoe.

Armed with only this simple implement, it was not practical to

create and maintain bottomland fields.

Forest farming depends almost entirely upon the aspect of

fire for success unless a sufficient labor bank is available to

clear and maintain the plot by other means. Since the direct

effect of fire in agricultural endeavors was to clear the

surface of refuse and kill most of the plants above the soil,

hillside plots were preferred over narrow floodplain plots. The

increased fuel moisture along the bottomlands would have

greatly reduced the fire’s intensity resulting in ineffective

burns and therefore more competition with the planted crops.

In contrast, the use of fire on hillside plots would have been

much more effective for several reasons. Among these are

the greatly reduced moisture tolerant understory and a faster

curing rate for the one hour and 10 hour fuels. Even with

comparable vegetation types within floodplain and hillside

plots, the fire will usually be more intense on a slope due to

increased heat transfer through convection and radiation to

the fuels ahead of the flame front.

Slash and burn agriculture along the Cumberland

Escarpment, which had its beginnings over 3000 years ago

was a common practice for farming the hill slopes well into

the current century (Otto 1983). During the 19th century

forest farming was the major cause of wildfires that burned

millions of acres of Kentucky’s timberlands. In 1880, for

example, 10 documented fires burned over 556,000 acres of

woodlands in the state. Nearly half of these fires were the

result of land clearing exercises (Sargent 1884:491).

It is somewhat paradoxal that while most primitive societies

claim a preference for meat, their diet for the most part

consisted of plants. As previously noted, Jones’ (1936)

analysis of the Newt Kash human fecal remains indicated

that the inhabitants were chiefly vegetarian. The food supply

of these people came primarily from cultivated plants and

those gathered from the wild. A similar pattern was reported

by Watson (1974) from the fecal data of the Mammoth Cave

area where over 50 per cent of the diet were products of

horticulture. Thus it seems that by the close of the first

millennium B.C. domesticated plants were a very important

component of the prehistoric diet. The use of broadcast fire,

an essential agent for forest clearing, especially in upland

environments of the Cumberland Plateau for agricultural

endeavors resulted in a transformation of the forest toward

more fire tolerant species.

Palynological Evidence

On of the oldest and still most effective methods used for

environmental reconstruction is the retrieval, identification

and counting of fossil pollen grains (Butzer 1982:173). By

knowing which plants were present and their relative

quantities, it is possible to draw conclusions about the

climatic and environmental conditions prevailing at the time

of deposition (Faegri and Iversen 1989). Quantification of

charcoal particles by both size and quantity can be employed

as an indicator of past fire histories. For example, charcoal

fragments less than 10mm in diameter are considered as

indicators of regional fires, whereas those greater than

50mm are assumed to have been generated from forest fires

within the immediate vicinity.

Cliff Palace Pond

Cliff Palace Pond is a small woodland pond, perched along

the crest of north-south oriented narrow ridge at 424 m

elevation in northeastern Jackson County, southeastern

Kentucky. This narrow ridge, known locally as Keener Point,

is ringed by a high, nearly continuous sandstone cliffline. The

sandstone cap is underlain by calcareous shales and

limestone that form the lower slopes and stream valleys.

Today, the dry ridgetop forest cover is composed by pines

(Pinus rigida, P. virginiana and P. echinata) intermixed with

scarlet and chestnut oak (Quercus coccinea and Q. prinus

L.). During the historic era chestnut (Castanea dentata)

occupied the ridgetops on the sandstone derived soils but

was absent on calcareous soils (Braun 1950). Mixed

mesophytic forests occupied the mesic slopes on calcareous

soils below the sandstone cliffs. Since the pond is located

within an important center of early-prehistoric plant

domestication the deposits offered an opportunity to examine

the relationship between prehistoric Native America use of

fire and plant production that was retrieved from the

surrounding archaeological sites.

The pond was cored in October 1996 resulting in the

recovery of a 142-cm sequence of Holocene deposits. The

sediment sequence was placed in firm synchronic age

classes based on well dated Holocene pollen diagrams

elsewhere (Delcourt and Delcourt 1987, 1997, 1998 and

Delcourt and others 1998). Charcoal particles, recorded by

size class, were also tabulated for each of the distinctive

pollen zones. Pollen diagrams were calculated as

percentages, representing fire-tolerant trees and shrubs

(figure 3) and fire-intolerant trees and shrubs (figure 4)

based on the sm totals of pollen grains and spores identified

in the sediment core. Figure 5 represents the charcoal

diagram in comparison with the major plant taxa represented

in the core sample. The following is a brief paleoecological

interpretation for each of the distinctive pollen zones.

Interval 9500 to 7300 Years Before Present.

During the early Holocene interval the forests near Cliff

Palace Pond were composed of cool-temperate to boreal

trees including spruce (probably red spruce, Picea rebens),

which made up nearly 5 percent of the forest composition.

The vegetation was dominated by cedar thought to have

been northern white cedar (Thuga occidentalis). Cedar

pollen, which reaches a maximum of 68 percent within this

interval would have been favored by the highly seasonal

climate of cold winters and warm summers. Hornbeam

(Ostrya/Carpinus type) alder (Alnus rugosa type) and birch
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 41

Figure 3.—Cliff Palace Pond pollen diagram,

fire-tolerant trees and shrubs. From Delcourt

and others 1998.

Figure 4.—Cliff Palace Pond pollen diagram,

fire-intolerant trees and shrubs. From

Delcourt and others 1998.
42 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

(Betula) also grew in abundance within the vicinity of Cliff

Palace Pond. Pines are represented by up to 2 percent and

oaks increase from 10 percent up to 35 percent within this

pollen zone. Pollen sums for fire-tolerant trees and shrubs

during this interval range from 7 to 20 percent with fireintolerant species comprising between 19 to 38 percent of

the assemblage.

During this period the native people are becoming less

nomadic and more territorial. Although they tended to live for

longer periods of time in one place, they were still nomadic

peoples, never staying in one place longer than a few

months. Their camps were placed within areas rich in a

variety of natural resources. Anthropogenic fires were used

primarily for game drives.

Interval 7300 to 4800 Years Before Present.

The Middle Holocene climate was warm but with lessened

extremes in temperature. The shifting storm patterns

associated with the “Hypsithermal Interval” resulted in

increased precipitation for southeastern Kentucky although it

was a time of warmth and drought in the midwestern portions

of the U.S. Within the vicinity of Cliff Palace Pond the

northern white cedar and spruce became locally extinct and

was replaced by the immigration and establishment of

temperate trees creating a mixed mesophytic forest

community including hemlock (1 to 6 percent), ash (up to 5

percent), basswood (1 percent) sugar maple (up to 4

percent), butternut (1 to 2 percent) and beech (trace

amounts). Oaks increase to 45 percent while pines remains

at less than 5 percent.

Although this is a time of increased moisture, the total values

for fire-tolerant taxa range from 8 to 25 percent. The range of

fire-intolerant taxa with a range between 16 to 31 percent

shows a slight decrease. These differences are thought to

reflect the steady increasing Native American populations’

continued use of fire to promote hunting and foraging.

Interval 4800 to 3000 years Before Present.

This interval begins with a major decline in the hemlock

composition of the forest community. Throughout its range in

eastern North America a widespread dieback of hemlock has

been observed around 4800 years ago. This is thought to

have been the result of an infestation of the hemlock looper

(Davis 1981). Coincident with the hemlock decline is a large

peak in the charcoal accumulation rates at Cliff Palace Pond

indicating a major fire event. Following the hemlock decline

and the catastrophic fire, extensive stands of eastern red

cedar (Juniperus virginiana) were established on Keener

Point. Cedar pollen dominates the pollen assemblage at 80

percent, with oaks persisting at 3 to 6 percent and ahs at 5

percent. Fire-tolerant trees and shrubs decrease to a

minimum of 2 percent, whereas fire-intolerant taxa increase

to a maximum of 82 percent.

Figure 5.—Cliff Palace Pond charcoal diagram and major plants represented by pollen grains and spores in

sediment core. The charcoal particles are tabulated in size classes of cross-sectional area, ranging from 0.5

grid to >5.0 grids (1 square grid = 165.12 μm2). After Delcourt and others 1998.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 43

Interval 3000 to 200 years Before Present.

Cedar declined dramatically in importance falling from 60

percent to trace amounts. Its demise was followed briefly by

a succession of ash (increasing from 6 t 36 percent, then

dropping to 5 percent). Subsequently, the forests of Keener

Point were dominated by oaks, rising from 11 to 55 percent;

chestnut, from 1 to 19 percent; and pine from 1 to 17

percent. During this interval pollen grains of both cultigens

within the “eastern agricultural complex” and weedy or

ruderal species such as ragweed that invade forest openings

made by humans activities appear within the record.

Paradoxically, during the Late Holocence time of climatic

cooling and increased participation, the forest on and around

Keener Point was dominated by fire-adapted taxa. The

overall values for fire-tolerant woody plants increase up to 82

percent, while fire-intolerant trees and shrubs diminish to

less tan 5 percent. This complete shift in forest composition

is considered to have been brought about in large part to

Native Americans bringing several plants under

domestication. The charcoal record for this interval shows a

steady increase in large charcoal particles indicating local

fires occurred within the immediate vicinity of Cliff Palace

Pond. This is in line with the slash and burn agriculture

techniques that would have been employed by the

prehistoric framers to clear their upland garden plots.

Interval A.D. 1800 to 1996.

During the last 200 years, a series of changes in forest

composition occurred following the total replacement of

Native American populations by the EuroAmerican

settlement of eastern Kentucky. The chestnut blight of the

1930s results in almost total eradication of this tree. This is

offset by an increase for oaks (54 to 58 percent), red maple

(7 percent) and black gum (3 percent). Total pine pollen

remains at about 10 percent but Virginia pine (Pinus

virginiana) increases locally. Following deforestation and

conversion of large portions of the landscape to agriculture

ragweed increases to 12 percent and exotic species

introduced from Europe such as plantain, dock and purslane

(Portulaca) are found within the pollen record for the first time.

For the first 100 years of this period, fire activity increased as

forested lands were cleared and burned for new agricultural

fields. Slash and burn agriculture, practiced by the American

Indians continued only on a larger scale. During the last 100

years fire prevention programs have dramatically reduced

large-scale fire occurrences.

Conclusion

As previously noted, the reconstruction of vegetational

histories must come from an interdisciplinary approach

between natural and cultural resource studies. Historic

accounts for example, can offer a multitude of facts about an

area with regards to the vegetative cover, fire occurrences,

climate, wildlife, land-use practices, etc. but within the New

World can only cover the last 500 years. Archaeological sites

often contain enormous amounts of ethnobotanical remains

that provide insights into the selection and utilization of

economically important plants over several millennia. These

can range from the selection of firewood and construction

materials to the propagation of plant food resources.

However, archaeological sites often lack the spatial

framework necessary to extract a record of how the human

groups affected their environments beyond the immediate

vicinity of the site. Pollen and charcoal analysis from

geologically deposited sediments on the other hand is a very

effective technique for reconstructing the vegetational and

fire history of a broader area but in most cases cannot

determine the causal factors for the change. Each of these

approaches provides a unique dimension to reading the

landscape. It is only through a multidisciplinary approach that

the tangible effects of fire on the landscape can be viewed

within a diachronic perspective.

The interaction of prehistoric human activities and forest

dynamics on the landscape took place by the way of the

interrelationships among the cultural use of fire, cultivation of

plants and forest succession. The changes upon the

landscape for the most part, occurred very gradually over

hundreds or thousands of years. The transition from on

forest type to another was so gradual that for those who

didn’t keep diaries, logs, or oral traditions, the change was

barely noticed. It is through archaeological and palynological

investigations that the murky waters of past forest conditions

become clear.

Through the combination of archaeological data from sites

within the general vicinity with the pollen and charcoal

analysis of Cliff Palace Pond, a link between forest

composition, fire history and the interaction of people with

their environment is beginning to emerge. The strong

coincidence of a temporal link between the domestication of

native plants, the prehistoric human use of rockshelters and

the increase of local fires as evident from the charcoal record

of Cliff Palace Pond argues for a cause-and-effect

relationship between Native American activities and changes

in forest composition. Within the Escarpment Zone of the

Cumberland Plateau prehistoric Native Americans began to

concentrate their activities in and near rockshelters on the

upper slopes and on ridgetops (Cowan 1985a,b; Ison 1988).

The use of fire especially in the preparation of the garden

plots adjacent to these campsites resulted in a fine-grained

patchwork of vegetation that included fire-adapted and fire

tolerant species on the upper slopes and ridgetops while still

allowing for the persistence of mixed mesophytic forest

communities on the lower slopes.

It should be quite clear that environmental reconstructions

cannot be accomplished without including people into the

equation. That fire undoubtedly shaped the ecosystems of

the southern Appalachians is evidenced by numerous

examples. The historical, archaeological and palynological

evidence from the Cumberland Escarpment has provided us

with the data we have for so long desired. It is now up to

land managers to put this data to work. Given the

documented fire history for the area we must once again

allow fire to play its natural role. If we do not, we will be the

progenitors of a forest that has never existed on the

landscape before.
44 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

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46 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Abstract

Oak distribution and dominance has exhibited major

changes since European settlement during the 18th and 19th

centuries. Large-scale increases in oak species have

occurred as a result of fire exclusion in the central tallgrass

prairie and savanna regions, cutting and burning of the

northern conifer and hardwood forests of New England and

the Lake States region, and land clearing, the charcoal iron

industry, and the eradication of Castanea dentata following in

the mid-Atlantic region. Studies of the dendroecology and

successional dynamics of several old-growth forests indicate

that prior to European settlement oak grew and regenerated

in uneven-aged conditions. At times oak growth was very

slow (< 1.0 mm/year) for long periods, which is usually

characteristic of highly shade tolerant species. Oak species

exhibited continuous recruitment into the canopy during the

17th, 18th and 19th centuries, but stopped recruiting in the

early 20th century. Since that time, later successional,

mixed-mesophytic species have dominated understory and

canopy recruitment, which coincides with the period of fire

exclusion throughout much of the eastern biome. Major oak

replacement species include Acer rubrum, A. saccharum,

Nyssa sylvatica and others. Logging of oak forests that have

understories dominated by later successional species often

accelerates the oak replacement process.

Basal cross sections were obtained from a partial timber cut

in two old-growth forest stands in Western Maryland, USA, in

1986. We recorded 42 fires from 1615 to 1958, which

occurred on average every eight years. There were no

significant differences in mean return interval between preEuropean settlement (before 1750) and the post-settlement

(1750-1900) periods. However, there were no major fire

years (25% of samples with scars) after 1900. Oaks

recruited consistently from the early 1600s to the early

1900s, but there was increasing amounts of A. rubrum and

B. lenta after 1900. Tree-ring dynamics, species recruitment

patterns, and long-term fire history reviewed in this paper

offer important direct support for the hypothesis that periodic

fire played an important role in the historical development of

oak forests of the mid-Atlantic region before and after

European settlement, and that fire suppression during the

20th century is facilitating a replacement of canopy oak by

later successional tree species.

Introduction

Oak species have a long history of domination in eastern

North America, and their present distribution in various

regions exceeds that recorded in the original forests prior to

European settlement (Abrams 1992). Much of the increase in

oak during the late 18th and 19th centuries can be attributed

to historical changes in disturbance regimes in the eastern

biome. Moreover, much of the expansion of oak has

occurred on xeric or nutrient-poor sites, which indicates the

stress tolerance capabilities of many oak species. However,

recent evidence indicates that oak forests, in the absence of

periodic understory fire, are often transitional to later

successional forest types. The purpose of this paper is to

review to linkages in the dynamics for oak species of the

eastern United States in relation to the historical changes in

land-use and fire regimes.

Historical Ecology of Eastern Oak Forests

Central Plains Example

Prior to European settlement, tallgrass prairie and oak

savannas dominated vast areas of the Central Plains,

southern Lake States and mid-western regions of the United

States (Kuchler 1964, Nuzzo 1986). Much of this region is

now part of oak-hickory forest association. Tallgrass prairie

and oak savannas in this drought-prone region were

maintained by frequent fire at 1-10 year intervals that were

initiated by Indian (Native American) activity or lightning

strikes (Cottam 1949, Day 1953, Pyne 1983, Abrams 1992).

Eastern Kansas represents the western limit of the eastern

deciduous forest, and oak species often grow along streams

and ravines forming relatively thin bands of “gallery” forests.

A study of the forests in a Kansas (Konza) tallgrass prairie

was completed to characterize the composition, structure,

development and successional dynamics of this oakdominated forest type (Abrams 1986). The range of sites on

Konza Prairie included mesic riparian benches to xeric

limestone ridges. Tree species importance varied with site

moisture relations with Celtis occidentalis - Q. macrocarpa,

Q. macrocarpa, Q. muehlenbergii - Q. macrocarpa, and Q.

muehlenbergii dominating forests along a continuum from

mesic to xeric, respectively. In each of the gallery forests

studied, oak species represented the oldest and largest

individuals, whereas the understory trees and regeneration

layers were dominated primarily by C. occidentalis, Ulmus

rubra and U. americana, and Cercis canadensis. An analysis

of the historical records, including the original land survey in

1858 and aerial photographs taken in 1939 and 1978,

indicated that the extent of the gallery forests has greatly

expanded.

This study exemplifies a major developmental pathway of

oak forests in the Central Plains. High fire frequency and

intensity during the period of Indian habitation maintained

tallgrass prairie species and retarded oak distribution,

relegating oak species to savannas and protected

woodlands. Following European settlement, the influence of

fire decreased due to road construction, expansion of towns,

cattle grazing, fire suppression activities, and the elimination

Fire and the Ecological History of Oak Forests in the Eastern United States

Marc D. Abrams1

1Professor of Forest Ecology, The Pennsylvania State University,

School of Forest Resources, 4 Ferguson Building, University

Park, PA 16802
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 47

of Indian fire activity (Pyne 1983, Abrams 1986). With less

fire, oak species expanded into the tallgrass prairie

vegetation, with Q. macrocarpa and Q. muehlenbergii

dominating mesic and xeric sites, respectively, in this

example. Thus, a significant proportion of the oak-hickory

forest in the former tallgrass prairie region is a recent

phenomenon in response to fire exclusion following

European settlement (Gleason 1913, Kucera 1960).

Lake States Example

Presettlement forests of the upper Lake States and northeast

were dominated by Tsuga canadensis, Pinus strobus, A.

saccharum, F. grandifolia and Betula allegheniensis, with

generally a very small percentage of Quercus (e.g., Q. alba,

Q. rubra and Q. velutina) (McIntosh 1962, Siccama 1971,

Finley 1976, Whitney 1986). In contrast, Quercus species

now represent a significant proportion of northern hardwoodconifer forests, and in particular Q. rubra has developed

prominence (Crow 1988, Whitney and Davis 1986). We

studied the presettlement forest records and current forest

composition and structure of 46 Q. rubra forests along an

edaphic gradient in north-central Wisconsin to gain an

understanding of their historical development and current

and future ecological status (Nowacki and others 1990).

Prior to European settlement, forests on mesic and

transitional mesic sites in the study area were dominated by

Tsuga canadensis, Betula, Acer and Pinus . Dry mesic sites

formerly comprised Pinus, Quercus (Q. velutina, Q.

macrocarpa and Q. alba) and Populus, while dry-mesic sites

were dominated by Pinus, Populus and Betula. In contrast,

many forests of the region are presently dominated by Q.

rubra, with relative importance values of 37-51% (Nowacki

and others 1990). Other important overstory trees included

Acer rubrum on transitional and dry mesic sites, A.

saccharum on mesic and transitional mesic sites, Q. alba on

transitional dry-mesic sites and Betula papyrifera on drymesic sites. Understory trees and reproduction layers were

dominated primarily by A. saccharum on mesic sites, A.

saccharum and A. rubrum on transitional sites, and A.

rubrum on dry-mesic sites.

The results of this study indicate that Q. rubra had a marked

expansion in northern hardwood-conifer forests following

Euro-American settlement. Quercus rubra on mesic and

transitional mesic sites developed following disturbance to

the original conifer-northern hardwood forests. Forests on

transitional dry mesic and dry mesic sites developed from

former oak-pine and pine forests, respectively. A postsettlement increase in Q. rubra has been documented in

other forests in the northeastern and Lake States regions

(Whitney 1986, Whitney and Davis 1986, Crow 1988,

Abrams 1992), and appears to be a direct result of

widespread cutting and subsequent fire in the middle to late

1800s and early 1900s. Evidence indicates that Q. rubra in

the overstory was present in relatively low numbers in

presettlement forest, but may have been pervasive in the

understory of the former pine forests. This coupled with the

widespread dispersal of acorns by birds and small mammals

facilitated the expansion of this species following large-scale

disturbances of the original northern hardwood-conifer

forests (Crow 1988).

Mid-Atlantic Region Example

Presettlement forests of southern New England and the midAtlantic region were dominated by Quercus in combination

with other species (Table 1). The leading tree species were

Q. alba, Q. velutina, Q. rubra, Q. pinus, Carya spp.,

Castanea dentata and Pinus spp. (including P. strobus, P.

rigida, and P. echinata). Evidence from eye witness accounts

and charcoal studies indicate that precolonial fires from

Indian activity and lightning strikes were pervasive in the

Table 1.—Presettlement forest types in the oak-hickory and oak-pine associations

of the eastern United States

Presettlement composition State References

Q. alba - Q. velutina - Pinus MA Whitney and Davis 1986

Q. alba - Q. velutina - Carya NY Glitzenstein and others 1990

Q. rubra - Castanea dentata NJ Ehrenfeld 1982

Q. alba - Q. velutina - C. dentata - Carya NJ Ehrenfeld 1982

Q. alba - Q. velutina - Carya NJ Russell 1981

Q. alba - Q. velutina - Carya PA Abrams and Downs 1990

Q. velutina - Q. alba - C. dentata - Carya PA Mikan and others 1994

Q. prinus - Q. alba - Pinus rigida - C. dentata PA Nowacki and Abrams 1992

Q. alba - P. strobus - Carya PA Nowacki and Abrams 1992

Q. alba - Q. rubra VA Spurr 1951

Q. alba - Q. rubra - Q. prinus - Carya - Pinus VA Spurr 1951

Q. alba - Q. rubra VA Orwig and Abrams 1994

C. dentata - Q. rubra VA Braun 1950

Q. rubra - Q. prinus - C. dentata VA Braun 1950, Stephenson 1986

Q. alba - C. dentata - Carya - Pinus WV Abrams and others 1995
48 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

region and probably played an important role in the

long-term stability of these forest types (Day 1953,

Watts 1980, Lorimer 1985, Patterson and

Sassaman 1988, Abrams 1992).

As in other regions of eastern North America,

disturbances associated with European settlement

had a dramatic impact on the original oak-hickory

and oak-pine forests. Widespread logging and

increased fire associated with land clearing, the

charcoal iron industry, tanbark and chemical wood

cuts, and lumbering of high quality hardwood and

conifers (e.g., P. strobus and Tsuga canadensis)

occurred during the initial settlement period (Pearse

1876, Abrams and Nowacki 1992, Russell and

others 1993, Mikan and others 1994). In one

example from central Pennsylvania, there were

nine active iron furnaces and ten forges in Centre

County in 1826, which were responsible for the

clearing vast forest acreage each year for charcoal

production (Abrams and Nowacki 1992). This type

of disturbance regime was responsible for

significant changes in species assemblages. In

central Pennsylvania, the original Q. alba - P.

strobus - Carya forests that were clearcut and

burned in the 1800s became dominated almost

exclusively by Q. alba and Q. velutina (Abrams and

Nowacki 1992). Cutting for charcoal in New Jersey

resulted in the increased dominance of Quercus

and Betula, and decreased dominance of Tsuga

and Fagus (Russell 1980). Quercus rubra

importance increased from 7% in presettlement P.

strobus forests in Massachusetts to nearly 20% in

present-day forests in response to landclearing and

logging (Whitney and Davis 1986). The decrease in

T. canadensis and P. strobus in these examples

can be related, at least in part, to their inability to

reproduce vegetatively.

Another major anthropogenic influence to eastern Quercus

forests has been the introduction of the chestnut blight

fungus (Endothia parasitica) during the early 1900s. This

fungus has been responsible for the elimination of overstory

C. dentata thought the eastern biome. The changes to

former chestnut dominated forests has been the subject of

several studies, most of which indicate that Quercus species

were one of the major beneficiaries of this disturbance. For

example, former oak-chestnut forests in North Carolina

became dominated by Q. rubra, Q. prinus, Q. alba and

Carya spp. (Keever 1953) (Tables 1 and 2). In southwestern

Virginia, Q. rubra represented 69% importance in forests

where C. dentata formerly comprised up to 85% of the

canopy (Stephenson 1986). In the ridges of central

Pennsylvania, Q. prinus, Q. rubra and Acer rubrum

increased where Castanea and Pinus were previously

important (Nowacki and Abrams 1992). Thus, postsettlement disturbances to eastern forests via land clearing,

the charcoal iron industry, lumbering and the chestnut blight

have led to increases in Quercus above levels estimated in

the original forest.

Q. alba and P. strobus dominated the original forests on

mesic valley floor sites within the eastern Ridge and Valley

Province, which extends from southeastern New York to

southern Tennessee (Braun 1950). The composition,

diameter and age structure, and radial growth chronologies

were studied in one of the few remaining undisturbed

remnants of this forest type located in southern West Virginia

(Abrams and others 1995). The forest is presently dominated

by P. strobus (34%), Q. alba, Q. rubra and Q. velutina (26%

total) and Acer rubrum (24%), and is uneven-aged with Q.

alba (max. age = 295 years) and P. strobus (max. age = 231

years) representing the oldest and largest trees (Fig 1). Q.

alba exhibited continuous recruitment into the tree size

classes from 1700-1900, whereas peak recruitment of P.

strobus occurred between 1830-1900. Interestingly, the

increase in P. strobus was followed by a wave of Q. rubra

and Q. velutina recruitment, suggesting possible facilitation

of these red oaks by P. strobus (cf. Crow 1988, Abrams

1992). After 1900, Pinus and Quercus recruitment stopped,

while that of A. rubrum, A. saccharum, F. grandifolia and T.

canadensis greatly increased.

Figure 1.—Age-diameter data for all cored trees and mean ring width index

for white oak and white pine and in an old-growth white pine-mixed oak

forest in southern West Virginia (adapted from Abrams and others 1995).

PIST = Pinus strobus, QUAL = Quercus alba, QURU = Q. rubra, QUPR =

Q. prinus ACRU = Acer rubrum, ACSA = A. saccharum, TSCA = Tsuga

canadensis, FAGR = Fagus grandifolia, CAGL = Carya glabra, NYSY =

Nyssa sylvatica.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 49

Radial growth analysis of the four oldest Q. alba indicated a

series of releases between 1710-1740, 1800-1830 and

1900-1930, with low or decreasing growth in the interim and

most recent periods (Fig. 1). In the early 1800s, releases in

radial growth were associated with high P. strobus

recruitment, while releases in the early 1900s coincided with

episodic A. rubrum recruitment. Individual radial growth

chronologies for trees of various species and age classes

indicated a series of major and moderate releases every 20-

30 years throughout the forest (Abrams, et al. 1995). The

asynchronous nature of these releases suggest a series of

small-scale disturbances with localized impacts.

We found evidence of fire scars, soil charcoal and windthrow

throughout the forest, and believe that these disturbance

factors significantly influenced the ecology of this old-growth

forest. Quercus and Pinus perpetuated themselves during

the 1600s, 1700s, and 1800s, but not in the 1900s, despite

evidence of blowdown during this century. These data are

consistent with the fire exclusion hypothesis, which led to a

Table 2.—Overstory and understory composition reported for Quercus forests in the major vegetation associations

of the eastern United States

Overstory Understory State Reference

Oak-hickory

Q. macrocarpa - Celtis occidentalis C. occidentalis - Ulmus KS Abrams 1986

Q. muehlenbergii Ulmus - Cercis canadenis - Q. muehlenbergii KS Abrams 1986

Q. velutina - Q. alba Prunus serotina - Acer negundo - Ulmus WI McCune & Cottam 1985

Q. marilandica - Q. stellata Q. marilandica - Q. stellata OK Rice & Penfound 1959

Q. alba - Carya ovata A. saccharum IL Rodgers & Anderson 1989

Q. alba A. saccharum MO Pallardy and others 1988

Q. alba - Q. velutina A. rubrum - A. saccharum - Betula lenta NJ Little 1974

Q. alba - Fagus grandifolia A. rubrum - A. saccharum - P. serotina PA Abrams & Downs 1990

Q. alba - Q. velutina A. rubrum - P. serotina PA Abrams & Nowacki 1992

Q. prinus - Q. rubra A. rubrum - Q. prinus - B. lenta PA Nowacki & Abrams 1992

Q. prinus A. rubrum - B. lenta - B. allegheniensis PA Mikan and others 1994

Oak-pine

Q. alba - Liriodendron tulipifera Nyssa sylvatica - Cornus florida VA Orwig & Abrams 1994

Q. alba - Q. coccinea N. sylvatica - A. rubrum - C. florida VA Orwig & Abrams 1994

Q. alba - Q. prinus - Q. coccinea N. sylvatica - Carya - C. florida VA Farrell & Ware 1991

Q. rubra A. saccharum - A. rubrum VA Stephenson 1986

Q. alba A. rubrum NC Christensen 1977

Q. rubra - Q. prinus - Q. alba Q. rubra - A. rubrum - Carya NC Keever l953

Maple-beech-basswood

Q. alba - Q. velutina P. serotina - A. rubrum MI Dodge & Harman 1985

Q. macrocarpa - Q. alba A. saccharum - Fagus grandifolia OH Cho & Boerner 1991

Q. alba - Q. rubra - Q. velutina A. saccharum - F. grandifolia - A. rubrum OH McCarthy and others 1987

Q. alba - Q. velutina - Q. rubra A. saccharum - F. grandifolia IN Schmelz & Lindsey 1970

Q. rubra - Q. alba - Q. macrocarpa A. saccharum - Aesculus glabra IN Parker and others 1985

Northern hardwood - Conifer

Q. velutina - Q. rubra - Pinus strobus A. rubrum MA Lorimer l984

Q. prinus - Q. rubra A. rubrum - A. saccharum NY McIntosh 1972

Q. velutina - Q. alba A. rubrum MI Host and others 1987

Q. rubra - Q. alba A. saccharum - A. rubrum WI Nowacki and others 1990

Q. ellipsoidalis P. serotina - A. rubrum WI Reich and others 1990

Mixed-mesophytic

Q. alba - P. strobus A. rubrum - F. grandifolia - Tsuga canadensis WV Abrams and others 1994

Q. alba - Q. coccinea - Q. prinus A. rubrum - A. saccharum - F. grandifolia KY Muller & McComb 1986

Q. alba - Q. velutina A. rubrum - N. sylvatica KY Campbell 1987

Q. prinus - Q. alba Q. prinus - Q. alba TN Schmalzer 1988

Southern evergreen

P. palustris - P. elliotii - Q. laevis Q. laevis - Q. geminata - Carya floridana FL Myers 1985

F. grandifolia - Q. laurifolia - Magnolia Liquidambar styraciflua - N. sylvatica - Fagus GA Quarterman & Keever 1962
50 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

shift in tree recruitment from Quercus and Pinus to Acer,

Fagus and Tsuga. Without intensive management in the

future, including prescribed fire, we predict this forest will no

longer support a significant Quercus and Pinus component.

Dendroecology of Old-growth

Quercus Prinus

We identified an old-growth Q. prinus forest on a dry talus

slope with canopy trees up to 326 years old at the Detweiler

Run in central Pennsylvania (Ruffner and Abrams 1998).

Unlike the transitional nature of eastern oak forest on mesic

sites, this xeric forest may maintain stable oak populations

even in the absence of fire. In 1996, Q. prinus at the study

site represented 39% importance, while A. rubra, Betula

lenta, Nyssa sylvatica, and Q. rubra represented a combined

50% importance. Q. prinus represented 75% of the canopy

dominant trees, but less than 10% of the intermediate and

overtopped trees. Continuous recruitment of Q. prinus in this

forest occurred between 1660-1950 (Fig. 2). Peak

recruitment periods for Q. prinus occurred during the late

1700s and early 1800s, which coincided with a release in

radial growth during this period.

Ironically, this forest is located near major 18th and 19th

century charcoal iron activity, where adjacent forests were

logged on a 20-30 year rotation. The extreme talus slope

undoubtedly protected this forest from cutting during that

period. Frequent cutting and occasional burning of most

forests in the region promoted oak coppicing and checked

the advance of later successional species until the late

1800s when charcoal iron production ceased in central

Pennsylvania. Fire suppression activities and less forest

cutting during the 20th century promoted Acer, Betula, and

Nyssa dominance in area forests. Because the extreme

edaphic condition, we predict that Q. prinus will remain a

dominant on the site with lesser amounts of more shade

tolerant Acer, Nyssa and Betula species. Thus, oak climaxes

may exist in the eastern oak region on very dry and nutrient

poor sites, such as this study area.

Successional Status of Eastern Oak Forests

Despite their importance throughout the eastern forest

biome, there is a strong indication that many oak forests are

transitional to later successional associations. Table 2 lists

the principal overstory and understory composition in various

oak forest types. Acer rubrum and A. saccharum are the two

most important understory species, and they most likely

represent the major replacement species of oak throughout

much of the biome (cf. Lorimer 1984, 1989, Abrams 19992,

1998). However, west of the Acer range, Celtis occidentalis

and Cercis canadensis appear to be important oak

replacement species (Abrams 1986). In the mid-Atlantic

region, south of A. saccharum’s range, Nyssa sylvatica may

be a future overstory dominant in current oak forests (Ross

and others 1982, Farrell and Ware 1991, Orwig and Abrams

1994). Fagus grandifolia is occasionally noted as an

important understory or overstory in oak forests, and these

are generally limited to the maple-beech-basswood and

mixed mesophytic associations. Q. alba forests in

southwestern Pennsylvania and northern Virginia are

presently dominated by F. grandifolia, A. rubrum and

Liriodendron tulipifera (Abrams and Downs 1990, Abrams

and Copenheaver 2000), indicating that F. grandifolia may

play an important successional role in eastern oak forests as

they move from second-growth to old-growth condition.

A few studies predict long-term stability of oak species on

very extreme sites. Examples of this include, xeric Q.

marilandica and Q. stellata forests derived from former oak

savannas in Oklahoma, Q. marilandica - Q. velutina in xeric,

upland glades in Illinois, mix-oak forests on nutrient-poor

barrens sites in New Jersey, dry talus slopes and Coastal

Plain forests in the mid-Atlantic region (Little 1974, Adams

and Anderson 1980, Dooley and Collins 1984, Orwig and

Abrams 1994, Ruffner and Abrams 1998). While these

forests may in fact represent edaphic climaxes they may

alternatively be exhibiting slow rates of successional

Figure 2.—Age-diameter data for all cored trees and mean ring

width index for chestnut oak in an old-growth mixed-oak forest in

central Pennsylvania (adapted from Ruffner and Abrams, 1998).
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 51

replacement and thus not have long-term stability in the

absence of fire. Clearly, the rates of succession on mesic

oak sites greatly exceeds that on xeric or nutrient poor sites

(cf. Abrams 1992). We still know very little about the

successional dynamics of oak forests in the southeastern

U.S. In one such study, A. rubrum is predicted to form a

dominant part of the overstory in a mature Q. alba forest in

North Carolina (Christensen 1977, R. Peet personal

communication). However, oak forests located to the south

and west of the peak distribution of A. saccharum, A. rubrum,

F. grandifolia and P. serotina may be experiencing less

successional pressure then oak forests in the northeastern

and north-central U.S. and may have stable oak populations

even in the absence of fire. Nonetheless, red maple density

increased by more than 300% in the subcanopy of a mature

pine-oak forest in southern Arkansas between 1954 and

1992 (Cain and Shelton 1995).

Fire Adaptations

In an early opinion survey of the fire resistance of 22

northeastern tree species, oaks (Q. prinus, Q. velutina, Q.

alba and Q. coccinea) where rated in four of the top six

positions (Starker 1934). It was further determined that these

oak species had a much greater bark thickness to trunk

diameter ratio than several mixed-mesophytic species, such

as A. rubrum, P. serotina, and F. grandifolia (Spalt and

Reifsnyder 1962, Harmon 1984). Among oak species, a

ranking of increasing bark thickness and fire resistance was

reported as follows: Q. macrocarpa > Q. velutina > Q. alba >

Q. rubra (Lorimer 1985, Hengst and Dawson 1994). Fire may

also be beneficial to oaks, relative to other hardwood

species, because they have relatively high resistance to

rotting after scarring, deep rooting and vigorous sprouting

ability, and increased germination and survival on firecreated seedbeds with reduced litter layers.

We compared the fire history and tree ring dynamics of two

old-growth forest stands in Western Maryland (Shumway

and others in press). The stands were located on the sideslopes of a ridge system (Savage Mountain) but differed in

the amount of periglacial material present. Both stands are

dominated by Quercus rubra, Q. alba, Q. velutina, Q. prinus

and Acer rubrum. Basal cross sections were obtained from a

partial timber cut in 1986, which provided evidence of 42

fires from 1615 to 1958 (fig.3). Fires occurred on average

every eight years, and there were no significant differences

in mean return interval among pre-European settlement

(before 1750), post-settlement (1750-1900) and fire

suppression (after 1900) periods. However, there were no

major fire years (25% of samples with scars) after 1900.

Oaks recruited consistently from the early 1600s to the early

1900s, but there was increasing amounts of A. rubrum and

B. lenta after 1900. There was a significant degree of

correspondence between the occurrence of major fires and

recruitment of oak species in either stand. Moreover, there

was an increase in radial growth in oak species in a fouryear period following the vast majority of fires, and this

increase was largely due to increased latewood production

following fires. Tree-ring dynamics, species recruitment

patterns, and long-term fire history reported in this study

offer important direct support for the hypothesis that periodic

fire played an important role in the historical development

and perpetuation of oak forests of the mid-Atlantic region

before and after European settlement, and that fire

suppression during the 20th century is facilitating a

replacement of canopy oak by later successional tree

species (Abrams 1992, 1998).

There exists a substantial database of paleoecological

studies that support the oak and fire hypothesis. For

example, at Crawford Lake in southern Ontario, beechmaple forests converted to pine-oak forests in response to

Iroquois cultivation and burning (Clark and Royall 1995).

Repeated disturbances (including fire) at a series of sites in

north-central Massachusetts led to a decrease in northern

hardwoods and an increase in pine, oak and chestnut during

the past 1000 years (Foster and Zebryk 1993, Fuller and

others 1998). Large oak pollen percentages were associated

with continuous charcoal influx during the Holocene in the

Hudson Highlands of southeastern New York (MaenzaGmelch 1997). A paleoecological investigation in eastern

Tennessee revealed that the study area was dominated by

oak, pine and ragweed during the last 1500 years, and that

charcoal influx increased during the period with increasing

human populations (Delcourt and others 1986). These

studies suggest a relationship among oak dominance, fire,

climate, and human activity.

Direct evidence for the role of fire in oak forests comes from

additional studies using the dating of fire scars. Most of

these data have been reported for oak forests in the central

plains, at the western extreme of the eastern forest, or from

pine-oak forests in southern Ontario (Abrams 1985; Guyette

and Cutter 1991; Cutter and Guyette 1994; Guyette and

others 1995; Guyette and Dey 1995). The mean fire return

interval (the time between fires) in these disparate locations

is surprisingly similar and ranges from 2 to 24 years. Fire

frequency either increased or decreased during the initial

period following European settlement depending on the

stand and region. A study of fire scars in a mixed-oak forest

that originated after 1850 in southeastern Ohio revealed that

fire occurred every 1 to 24 years, and averaged 7.5 years

(Sutherland 1997). Two black oak (Quercus velutina Lam.)

woodlands in Indiana had a fire return interval of 5.2 and

11.1 years during the 1900s (Henderson and Long 1984).

Six fires scars dating from 1641 to 1711, with a mean return

interval of 14 years, were reported from a pre-European

settlement oak in Mettler’s Woods, New Jersey (Buell and

others 1954).

Conclusions

The rise in oak dominance in the eastern U.S. at the

beginning of the Holocene epoch (10,000 y. B.P.) was

associated with warmer and drier conditions and the

increased occurrence of fire (Watts 1980, Davis 1985, Webb

1988). It is well documented that American Indians actively

used fire for a multitude of purposes, and they were probably

responsible for increasing the incidence of forest and prairie

fires above that caused by lightning strikes (Gleason 1913,

Day 1953, Pyne 1985, Patterson and Sassaman 1988).
52 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

While fires were too frequent in the tallgrass prairie region

and too infrequent in the northern hardwood forests for oaks

to prosper, the intermediate frequency and intensity of fire in

presettlement oak-hickory, oak-chestnut and oak-pine forests

were apparently necessary for long-term oak stability

(Abrams 1992).

Initially following European settlement, oak populations

throughout much of the eastern biome increased due to fire

exclusion in tallgrass prairie and southern pine forests,

widespread logging and burning of northern hardwoodconifer forests, and logging, burning and the chestnut blight

in the eastern mixed-oak forest types. Moreover, oak species

have typically shown a strong affinity for drought-prone sites,

which are fairly common in the eastern U.S. from extreme

edaphic, physiographic and/or climatic factors. Despite their

low to moderate shade tolerance rating, eastern oak species

maintained themselves in pre-European settlement forests in

uneven-aged conditions, often growing very slowly for long

intervals. Recurring fire in presettlement oak understories

most likely prevented significant invasion by later

successional species. This coupled with adequate light

transmission through oak canopies probably facilitated oak

establishment and recruitment in presettlement forests.

Consistent with these ideas, oak species exhibit a suite of

adaptations for fire.

Widespread invasion of most oak understories, excluding the

most xeric and nutrient poor sites, by later successional tree

species and a lack of oak recruitment coincides with the start

of fire exclusion in the early 1900s. It has been argued that

continued fire exclusion will lead to a vast reduction in oak

dominance in the eastern forest (Lorimer 1985, Abrams

1992, 1998). However, white-tailed deer populations have

also risen dramatically in many eastern region during the

1900s. Considering that many oak species are highly

preferred browse species, large deer populations are

probably exacerbating the oak regeneration problem

(Hughes and Fahey 1991, Abrams 1998). Thus, the increase

in Acer and other later successional trees in many oak

forests may be a function of fire exclusion, differential deer

browsing, as well as other less well documented factors.

Therefore, without intensive management to reduce

competition from non-oak species and the predation of

seeds and seedlings, such as through the increased use of

fire and controlling deer populations, a major loss of

overstory oak dominance can be anticipated for the near

future.

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Sutherland, E.K. 1997. History of fire in a southern Ohio

second-growth mixed-oak forest. Gen. Tech. Rep. NC188. U.S. Department of Agriculture, Forest Service,

North Central Research Station: 172-183.

Watts, W.A. 1980. Late quaternary vegetation of the

central Appalachian and the New Jersey coastal plain.

Ecological Monographs. 49: 427-469.

Webb, T. III. 1988. Glacial and Holocene vegetation

history: eastern North America. In: Huntley, B.; Webb,

T., III, eds. Vegetation History. Kluwer Academic Publ,

Amsterdam, The Netherlands: 385-414.

Whitney, G.G. 1986. Relation of Michigan’s presettlement

pine forests to substrate and disturbance history.

Ecology. 67: 1548-1559.

Whitney, G.G. 1987. An ecological history of the Great

Lakes forest of Michigan. Journal of Ecology. 75: 667-

684

Whitney, G.G.; Davis, W.C. 1986. Thoreau and the forest

history of Concord, Massachusetts. Journal of Forest

History. 30: 70-81.

Woods, F.W.; Shanks, R.E. 1959. Natural replacement of

chestnut by other species in the Great Smoky

Mountains National Park. Ecology. 40: 349-361.
56 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Abstract

This paper reviews the scanty available database on the

effects of dormant season fire on ecological processes and

organisms in the forest floor and soils of forests of the central

hardwoods region. Data from studies in other ecosystem

types are used to supplement the existing database, but

extrapolation across ecosystem types should be viewed with

caution. Fires in the central hardwoods are typically low in

intensity and consume primarily the unconsolidated leaf litter.

As long as the fire can move across the open forest floor, soil

temperatures generally do not increase enough to cause

significant heating-induced mortality among soil-dwelling

organisms. Soils under smoldering piles of woody fuels may,

however, be subject to sterilization. Direct N volatilization is

probably not an important pathway for fire-related nutrient

loss due to low fire temperatures. The microclimate at the

forest floor surface is probably affected significantly and this

may produce phenological changes in root growth and

microbial activity. More research in this area is warranted.

Base cations released from dissolving ash may or may not

increase soil pH and cation availability, depending on the

nutrient status of the soil and the amount of ash deposited.

Nitrogen availability typically increases after one or a small

number of fires but may decrease over the long term.

Abundances of soil animals in the forest floor are reduced by

fire whereas those in the mineral soil are affected little.

Recolonization of the redeveloping post-fire forest floor is

rapid. Microbial abundances in the forest floor are typically

reduced by fire, but rapid recolonization by these groups is

also likely, except under smoldering piles of woody fuels.

The database for the central hardwoods region is scanty and

this area is greatly in need of additional research attention.

Based on this scanty database, belowground impacts of low

intensity, dormant season fires in the central hardwoods is

likely to be less that the impact aboveground, though

considerably more research covering a greater range of

ecosystem types and fire frequencies is needed to verify this.

Introduction

The history of ecological studies indicating the importance of

relatively frequent fire in the maintenance of forest

composition and structure in eastern North America goes

back over half a century. For example, Cottam (1949)

concluded that frequent fire played a key role in the

prehistoric ecology of oak-hickory (Quercus-Carya) forests in

Wisconsin, and Daubenmire (1936) emphasized the role of

fire in the vegetation of the “big woods” region of Minnesota.

Based on pollen and charcoal profiles, Delcourt and Delcourt

(1997) have determined that fire has been common in this

region for at least 4000 yr. Their analysis of the fire record

indicates, however, that the importance of local fires has

increased and that of regional fires has decreased over time.

The importance of local, relatively small fires in this region is

clear from the analysis of fire occurrence over the last

century in oak forests in Vinton County, Ohio done by

Sutherland (1997). In this region, the great majority of fires

occurs either during the dormant season (69%) or in early

spring (25%), with large, regional summer fires being rare

(Sutherland 1997). The mean fire interval for the Vinton

County sites was 3.6 yr, with locally extensive fires occurring

on the average of 7.5 yr (Sutherland 1997). Based on these

and other similar records of fire in the central hardwoods

region (e.g. Guyette and Cutter 1991), this remainder of this

review will focus on the effects of relatively low intensity,

dormant season fires on the ecology of the hardwood forest

floor and soil.

Forest fires may affect the belowground parts of the

ecosystem through direct heating and by consuming and/or

changing the characteristics of the surface fuels (Figure 1).

Such alterations to the ground surface may then affect

belowground organisms and ecological processes through

direct volatilization of N and S, microclimate alteration, and

deposition, dissolution, and convection of ash. In the

paragraphs that follow I attempt to review each of these

impact components individually, and conclude with a review

on the biological and ecological consequences of these

impact components taken together. One important caveat is

necessary, however, before proceeding: the direct database

for the central hardwoods appears to be sparse, at best. This

is particularly apparent when comparing the data from this

region to what is available for western conifer forests,

shrublands, and prairie (reviews by Wells et al. 1979 and

Neary et al. 1999). My approach is to summarize the data

from the central hardwoods and supplement where required

with studies from other temperate, humid ecosystems.

Fire Behavior and Direct Heat Effects

Low intensity dormant season fires rarely produce the large

flame fronts and extreme temperatures of dormant season

fires in chaparral or growing season fires in conifer forests

(Boerner 1982). Franklin et al. (1997) recorded surface fire

temperatures during two prescribed burns in forests near the

Kentucky-Tennessee border. They reported average

temperatures at the surface of the forest floor of 226C in an

oak-maple forest and 190C in a mixed oak site. Similarly,

Boerner et al. (2000b) reported mean fire temperatures of

210C and 157C in two early spring prescribed burns in oakhickory stands in southern Ohio, and Blankeship and Arthur

(1999a) reported forest floor surface temperature ranges of

316-398C and 205-315C in two prescribed fires in oak-pine

forests in Kentucky.

Effects of Fire on the Ecology of the Forest Floor and Soil

of Central Hardwood Forests

Ralph E.J. Boerner 1

1Professor and Chair, Department of Evolution, Ecology, and

Organismal Biology, Ohio State University, Columbus, OH

43210.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 57

The range of variability in forest floor surface

temperatures that can occur in this region is well

illustrated by the studies of prescribed fires in northern

Indiana forest stands conducted by Cole et al. (1992).

Low creeping fires in oak forests in their sites often

burned only leaf litter and produced little surface

temperature rise when burning downhill; in contrast,

peak temperatures 15 cm above the surface in other

stands reached 300-400C with very high spatial

variability (Cole et al. 1992).

Variations in fire intensity are related to many factors,

including forest floor biomass/depth, slope position,

aspect, and angle, and fire weather (Franklin et al.

1997; Boerner et al. 2000b). As an illustration of this,

Boerner et al. (2000b) compared the behavior of

prescribed fires done in two sites over three days in

which the weather changed from warm and relatively

dry to cool and moist. The fire conducted during the

relatively drier, warmer day attained average

temperatures at 10 cm above the forest floor of 210C,

with a strong variation in maximum fire temperature related

to slope position and soil moisture content. In contrast, the

fire conducted during cooler, moister conditions only attained

an average of 157C and there was no significant correlation

between maximum fire temperature and slope position or soil

moisture content (Boerner et al. 2000b). In the former fire, 2/

3 of sample plots attained maximum surface temperatures

>200C at least transiently, whereas in the latter fire only 1/6

of sample plots reached 200C (Boerner et al. 2000b).

Fires of this intensity typically burn only unconsolidated litter

and fine woody fuels, leaving the humus and upper soil

layers uncombusted. For example, in the fires described

above (Boerner et al. 2000b); litter loss from a single fire

ranged from <30% to >80%, depending on landscape

position and fire temperature, but the underlying humus layer

was unaffected. These results were similar to those obtained

in a series of late autumn prescribed burns in oak-pine

forests in southern New Jersey in which Boerner (1983)

observed losses of 50-70% of the litter mass but <5% of

humus mass, as well as an analysis of the impact of a single

winter fire on oak-pine forest in Kentucky which

demonstrated losses of ~30% of litter mass but no humus

(Blankenship and Arthur 1999b). Studies of fires in northern

Indiana oak forests and Georgia piedmont forests have also

indicated that the effects of direct combustion are typically

limited to the unconsolicated leaf litter (Cole et al. 1992;

Bender and Cooper 1968).

Although the underlying, humified portion of the forest floor

and the upper mineral soil layers are typically not combusted

directly in such fires, transfer of heat to the unburned forest

floor and soil could produce ecological consequences if

severe enough. For example, soil enzymes will denature

when temperatures reach 70C even transiently, and

temperatures >70C for 10 min will kill fungi, protozoa and

some bacteria (Lawrence 1956).

In general, however, surface fires moving across the open

forest floor do not have strong potential for severe soil

heating because only approximately 5% of heat released by

fire is partitioned to soil, and mineral soil is a very poor

conductor of heat (Raison 1979). Although there are few

studies that report mineral soil temperatures during fire in the

central hardwoods, there are studies from other regions that

can serve as examples of what one might actually expect in

our region. For example Saa et al. (1993) recorded mineral

soil temps at 5 cm depth were <50cm in fires in pine forests

and gorse (Ulex europaea) shrublands in Spain. Heyward

(1938) recorded soil temperatures during a fire with 1m

flame height in a North Carolina longleaf pine (Pinus

palustris) forest and found temperatures at 2.5cm into

mineral soil of 40C just after the flame front passed and only

25C 45 minutes after the fire had passed.

Such fires can, however, have significant effects on

temperatures within the forest floor layers that remain largely

unconsumed by fire. Blankenship and Arthur (1999a) report

temperatures as high as 315C in the upper 0.5 cm of the Oe

layer during a fire in a Kentucky oak-pine forest that

generated temperatures as high as 398C at the forest floor

surface. Similarly, Cole et al. (1992) measured temperatures

in the humus layer during prescribed fires and recorded

ranges of 100-133C at 2 cm depth into the humus and 40-

90C at 3 cm into the humus. Thus, low intensity fires moving

across the open forest floor do not appear to have the

potential to cause significant heating-induced mortality to

organisms living in the lower humus layers or upper mineral

soil, but may have such effects near the litter/humus

interface.

The one situation in which direct, heating-induced mortality

of soil organisms may become significant is when localized

concentrations of woody debris continue to smolder in place

for an extended period of time. Miller et al. (1955) monitored

ground surface and soil temperatures during and after a fire

in a New Zealand shrubland. The main fire area experienced

maximum surface temperatures of about 200C, but there

was no change in temperature recorded by sensors buried 5

Figure 1.—Schematic approach for analyzing the direct and indirect

effects of fire on organisms inhabiting the soil and forest floor.

Fire and the Hardwood Forest Floor

Direct Effects

of Heating

Biological and Ecological Consequences

Chemical and Biochemical

Consequences

Effects of Fuel

Consumption

Volatilization

of Nitrogen

Ash and Altered

Organic Matter

Microclimate

Alteration
58 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

cm and 10 cm into mineral soil. However, in areas where

localized wood heaps burned for several hours, sensors

recorded maximum temperatures of 100C at 5 cm and

60C at 10 cm depth in mineral soil. Thus, localized woody

fuel accumulations do present a situation in which direct

heating effects may have negative effects on soil biota.

Nitrogen Volatilization

Direct volatilization of N during combustion of litter can

result in large losses of N from the ecosystem. In a review

of the impacts of fire on nutrient cycling, Boerner (1982)

reported net losses of 30-100% of litter N during fire as

the result of direct volatilization and ash convection

(Figure 2). However, direct volatilization of N does not

begin until fire temperatures exceed 200C and does not

become a major pathway for N loss until temperatures

exceed 300C (Raison 1979; Boerner 1982). Given the

ranges of surface temperatures recorded in fires in the

central hardwoods noted earlier, it seems unlikely that

direct N volatilization will be significant in this region. For

example, Boerner et al. (2000b) estimated direct N

volatilization from two prescribed burns in Ohio oak-hickory

forests to be <1 kg/ha, less than 10% of annual additions to

the site through atmospheric N deposition.

In addition to losses due to direct volatilization, nutrients may

be lost from the forest floor via convection of ash (Boerner

1982). Monitoring of precipitation and ashfall in sites

adjacent to active fires in western conifer forests (Clayton

1976) and southeastern pine forests (Lewis 1974) have

demonstrated that nutrient redistribution via ash convection

and deposition can have significant effects on nutrient

budgets of neighboring sites.

Microclimate Alteration

Removal of the unconsolidated litter and deposition of

blackened, partially combusted material on the ground

surface have the potential to alter microclimate at the soil

surface. Although such effects have commonly been

reported in grassland ecosystems (Boerner 1982), to date

I’ve been able to locate no published microclimate data for

burned sites in the central hardwoods. However, in an

unpublished study from southern Ohio, early spring surface

soil temperatures in sites burned annually for three years

were higher during the day and lower during the night than

were those in soils of unburned controls (Personal

communication. 1999. Louis Iverson and Todd Hutchinson,

Northeastern Research Station, Delaware, OH 43015). Viro

(1974) presented air and soil temperature data for burned

spruce plantation in Finland. Soil temperatures were both

higher in the burned site than under the canopy in a

neighboring unburned site. However, soil temperatures in a

canopy gap in the unburned site were intermediate between

those of the burned site and those under the intact canopy.

Thus, in this site at least, both the opening of the canopy and

the alteration of the ground surface conditions contribute to

microclimate alteration.

What impacts might earlier warming and more extreme

conditions have on soil biota? In grasslands, the earlier

warming of soils in burned sites contributes to an increase in

the rate of N mineralization and nitrification, compared to

unburned sites (Boerner 1982). This may well occur in

central hardwoods sites as well but has not been explicitly

measured. However, Dress and Boerner (submitted)

observed that spring increases and summer decreases in

fine root biomass in an Ohio oak-hickory site burned three

times over three years were shifted 1-2 months earlier than

in unburned controls or sites burned once two years before.

The potential impact of ground surface microclimate

alteration on belowground ecology in the central hardwoods

remains largely unexplored.

Ash Deposition, Dissolution,

and Their Effects on Soil Chemistry

The material that remains after fire has consumed part or all

of the litter layer is a combination of partially-combusted

organic matter and inorganic ash. This inorganic ash is rich

in base cations such as Ca, Mg, and K, and is easily

dissolved by rainfall. Whether the dissolution of this ash has

subsequent effects upon surface soil chemistry depends, to

a great extent, on the nutrient status of the soil prior to the

fire and the mass of ash-derived nutrients that are added as

a consequence of fire.

The impact of ash deposition and dissolution can be most

vividly seen in studies of sites in which fire has been both

frequent and spatially concentrated. Mikan and Abrams

(1995) examined soil chemical properties in an oak forest

site in southeastern Pennsylvania that had been used as the

charcoal hearth for an iron plantation from 1771-1884. The

soils under the former hearths were 0.25 units higher in pH,

and base saturation increased from 10% in nonhearth soils

to 38% in hearth soils. Furthermore, exchangeable Ca was

12.1X higher and Mg was 4.8X higher in hearth soils than

Percent N Volatilized

0 20 40 60 80 100

Longleaf Pine, NC

Pine Plantation, SC

Mixed Conifers, WA

Chaparral, CA

Grasslands, UK

Heather Moorland, UK

Volatilization of N by Fire

(source: Boerner 1982)

Figure 2.—Proporation of fuel N content lost to direct volatilization

and ash convection. Original data sources are cited in Boerner

(1982).
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 59

nonhearth soils. Thus, the effects of repeated ash deposition

could still be observed even 110 years after the last hearth

fire.

A less extreme example comes from a study of the effects of

burning of concentrated slash piles after logging in an East

Anglia, England oak forest (Jalaluddin 1969). In this

situation, the slash was concentrated in 1.5-2.0m diameter

piles and burned, with active flames and smoldering lasting

>3hr. Samples taken several days after the fires indicated

that pH had increased from 6.0 in unburned to 9.0 under ash

piles. However, within six months, soil pH under the former

slash piles returned to preburn levels. Thus, the effects of

fire on soil chemical properties are sensitive to both the

amount of ash deposited in a fire and the number of fires that

occur at that site over time.

Eivasi and Bryan (1996) have studied the impact of longterm prescribed fire on southeastern Missouri oak-hickory

upland flatwoods. This experiment established in 1949, with

annual burns and periodic burns (every four years) done in

April prior to leaf-out. After 40+ years of burning there were

no significant effects of burning on soil pH, Ca, K, or Al, and

only a small indication of lowered Mg in annual burns.

However, available P was only 24% of control in the annual

plots and 35% of control in the periodic burn plots. Thor and

Nichols (1973) report similar results from an annual and

periodic burning study in the Highland Rim region of Franklin

County, Tennessee. After 8 annual or 2 periodic burns, there

were no statistically significant differences in pH among

treatments. DeSelm et al. (1991) studied the same

Tennessee site 27 years after the onset of burning. Again,

the pH and availability of K and P of soils in the burned plots

did not differ significantly from those of the controls. Thus, in

the loess/limestone-derived soils of these two sites, lengthy

periods of burning had no striking effect on base cations, but

did impact P availability negatively in site with the longer

history of burning.

Knighton (1977) studied the effects of 0, 1, 2, or 3 years of

annual burning on an oak-hickory forest in central Wisconsin.

Availability of Ca and Mg were considerably higher in the soil

solution from the burned sites than the control, but with no

indication of a cumulative effect over time. The soil solution

concentrations of NO3

-

 and PO4

3- were also greater in the

burned sites, and the average concentrations of both

appeared to increase with the number of fires. However,

Knighton (1977) points out that the increasing trend was not

statistically supportable due to high among-sample variability

in soil leachate samples. Blankenship and Arthur (1999b)

report an increase of 0.2-0.3 pH units and a transitory

increase in available N following a winter prescribed fire in

an oak-pine stand in eastern Kentucky. They did not,

however, report increases in extractable cations to

accompany the increase in pH.

In our continuing study of annual and periodic prescribed fire

in southern Ohio we have observed a somewhat different

pattern of pH change. Our experimental design is based on

four study sites, each of which is divided into three

watershed-size treatment units. One of the units within each

study area has been burned annually since 1996, a second

was burned in 1996 and 1999, and the third remains as an

unburned control. In 1995, prior to any burning, soil pH did

not vary significantly among the three watershed-scale units

in three of our four study areas (Figure 3A; Boerner,

unpublished data). In the fourth site (Watch Rock), one unit

had significantly greater pH than the other two, although we

have still not yet determined why this difference existed.

Following 2-4 fires, we now see a significant and positive

effect on soil pH among all four study areas pooled and

within two of the four individual sites (Figure 3B; Boerner,

unpublished data). One site (Young’s Branch) has not

responded in soil pH to the burning. At Watch Rock, the one

treatment unit with the unusually high pH has remained an

outlier; however, the other burned unit at Watch Rock is now

significantly greater in soil pH than its unburned control. Our

study sites differ from those studied by Eivasi and Bryan

(1996), Thor and Nichols (1973) and DeSelm et al. (1991) in

that the soils of our sites were formed on nutrient and base

poor sandstones and shales, not on relatively nutrient-rich

loess and limestone as in the other studies. Thus, the impact

of the base cations dissolved from the ash deposits is likely

to be greater and more easily resolved in a nutrient-poor site

such as ours than in the richer limestone and loess sites.

In a study of site preparation burning prior to restoration of

Appalachian pine-mixed hardwood forests in North Carolina,

Knoepp and Swank (1993) demonstrated that soil NH4

+

concentrations and N mineralization rates increased after

fire. The additional N made available from ash dissolution

and from enhanced organic matter mineralization did not,

however, result in increased stream water N concentrations;

thus, the increased available N remained within the

ecosystem.

Despite considerable combustion of surface organic matter

by fire, few studies have noted changes in soil organic

matter content. Knighton (1977) observed no effect of 1-3

fires on soil organic C in a Wisconsin oak-hickory site, and

we have observed only slight increases in soil organic C in

our Ohio oak-hickory sites (Boerner et al. 2000b); Boerner

and Brinkman, submitted): Even after 40+ yr of burning in a

Missouri oak-hickory forest organic C content in the mineral

soil did not differ from that of unburned controls (Eivasi and

Bryan 1996).

It should be emphasized again at this point that these

patterns of effects are specific to dormant season, low

intensity fires. The effects of high intensity wildfire may be

quite different. For example, soil organic C and N availability

were reduced by as much as half following intense wildfire in

Pinus halepensis-Quercus calliprinos forests in Israel (Kutiel

and Naveh 1987)

Biological and Ecological Effects

For convenience, I will review the biological and ecological

effects of fire on soil organisms in decreasing size order:

from roots to soil animals to microbes. The only study of the

effect of fire on root biomass and production in the central

hardwoods comes from our own study sites. Dress and
60 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Figure 3.—A-horizon mineral soil pH in forested watersheds in southern Ohio. N=18 for each histogram bar

(except for pooled data in B where N=72) with standard errors of the means shown. A. 1995 prefire

variations among watershed-scale treatment units within the four study areas. B. 1999 postfire variations

among treatment units. Histogram bars labelled with different lower case letters were significantly at p<0.05.

Boerner (in press) determined that both live and dead fine

root biomass were significantly lower in a site burned

annually for three years than in an unburned control. The

variations in root biomass were inversely proportional to

variations in soil N availability and N mineralization. This

observation was consistent with both general models of the

relationship between root biomass/production and N

availability (Chapin 1980) and with studies in grasslands

that also demonstrated reciprocal responses of roots and N

availability to fire (Benning and Seastedt 1997).

The assemblage of small animals in the forest floor and soil

of a forest ecosystems is both extremely diverse and

critically important. These animals form a complex food web

that is responsible for much of the processing of the detritus

produced both above ground and below ground. Soil

animals such as springtails (collembola), mites, and

millipedes are important in breaking down coarse organic

matter such as leaves into fine material that is suitable for

microbial colonization and decay. Without soil animals

present, leaf litter decomposition essentially ceases

(Edwards and Heath 1963)

Once again, the database for the central hardwoods is

sparse. In our study area, the abundance of oribatid mites in

the top 15 cm of soil and forest floor decreases with

increasing fire frequency (control, one burn, three annual

burns), but the abundance of other mite groups was lowered

only in the annual burn site (Dress, unpublished data). Metz

and Farrier (1971) monitored the abundance of mites and

collembola in the forest floor and soil of a North Carolina pine

forest. In the forest floor layers, abundance of both groups

decreased with increasing fire frequency. However, when fire

was applied only every 4 years there was sufficient time for

recolonization and population growth to near control levels.

Abundances of collembola and mites in the mineral soil were

not affected by either annual or periodic burns in these sites.

Thus, on the basis of a scanty database we suggest that

animals in the litter are likely to be more affected than those

in the mineral soil and that effective repopulation can occur

under a periodic but not annual burning regime.

The effects of fire on the soil microbial community can be

evaluated by examining directly the effects on individual

species or species groups or by determining the effect of fire

on microbially-mediated processes such as N mineralization.

Jorgensen and Hodges (1970) analyzed the impact of annual

and periodic winter burns on the microbial community

structure of a loblolly pine (Pinus taeda) forest on the South

Carolina piedmont. In this study, the last periodic burn was
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 61

eight years earlier and last annual burn was slightly less than

one year earlier. In the mineral soil, none of the burning

treatments significantly affected the abundance of either

fungi or bacteria+actinomycetes, nor did burning affect the

diversity of fungi in either the mineral soil or the forest floor.

In the forest floor of the annual burn plots, however, the total

abundance of fungi was only 78% of that in the control and

the total number of bacteria+actinomycetes was only 18% of

the control level. Thus, the microbial abundance in forest

floor was strongly reduced by the annual burning. In

contrast, microbial abundance in the periodic burn site was

greater than in control, by 2.2X for fungi and by 1.2X for

bacteria+actinomycetes. So recovery from burn happened in

<8yrs and actually produced a forest floor environment more

suited to microbial activity than the unburned control.

The potential of fungi present in the mineral soil to facilitate

recolonization as the forest floor redevelops after fire is also

supported by the work of Tresner et al. (1950) in hardwood

forests of southern Wisconsin. They noted that although

numbers of fungi decrease from forest floor down to mineral

soil, and with depth in mineral soil, species composition does

not change, so colonization of disturbed surface layers from

deeper soil is possible without changing community structure

(Tresner et al. 1950).

A more extreme example of the impact of fire on fungi comes

from the work of Jalaluddin (1969) on the microbial ecology

of soils under small plots on which concentrated piles of

slash had burned and smoldered had continued >3hrs. At

one week, three months, and six months post-fire, fungal

abundance in the center of the former ash piles were only

3%, 6%, and 11% of that in areas >3m from fire piles. At the

edge of the burned areas where mycelial recolonization

would have supplemented colonization from new spores,

one week, three month, and six month abundances were

17%, 31%, and 43% of those >3 m from the burned areas.

Thus both mycelial ingrowth from areas surrounding the fire

and new spore colonization occur. However, in this case,

recolonization from below was prevented by the impact of

the lengthy smoldering of the fire sterilizing the lower soil

layers.

Mycorrhizal fungi are key symbionts for virtually all forest

plants. There are two major groups of mycorrhizal fungi

found in this region. The ectomycorrhizal fungi are a group of

higher fungi (basidiomycetes and ascomycetes) which form

symbioses with conifers, oaks, hickories, and beech. The

arbuscular mycorrhizal fungi are a group of lower fungi

(zygomycetes) that form symbioses with herbaceous plants

and woody plants other than those that depend on

ectomycorrhizae. In western conifer forests, some

ectomycorrhizal fungi seem to be sensitive to fire while

others are unaffected (Schoenberger and Perry 1982). Thus,

in western conifer forests, how seedlings of a host tree

species perform in recently burned sites may depend on

which fungi that species depends upon to form

ectomycorrhizae. Unfortunately, little is known about either

the diversity of either group of mycorrhizal fungi in eastern

forests or the response of these organisms to fire.

There may also be an interaction between effects on soil

animals and effects on microbes. Lussenhop and Wicklow

(1984) determined that fungal species diversity in Wisconsin

prairie plots subjected to spring burning and raking in a

Wisconsin prairie increased by 29% compared to the control.

Moreover, the fungal propagules were also less aggregated

and more evenly distributed in the burned plots. Lussenhop

and Wicklow (1984) feel this change in spatial dispersion

was due more to the increase in the abundance of mites and

collembola on the burn plots than the actual burning and

raking treatment because changes in fungal spatial

distribution correlated most closely with changes in fauna.

Other than the study of Jorgenson and Hodges (1970)

described above, studies of the direct impact of fire on

bacterial abundances and/or community composition in the

central hardwoods are uncommon. Blankenship and Arthur

(1999b) did find a significant, positive effect of a single

prescribed winter burn in an eastern Kentucky oak-pine

stand on bacterial biomass and also reported a decrease in

the fungal:bacterial biomass ratio following fire. Clearly the

impact of fire on the forest floor and soil microbial

assemblage continues to be neglected in fire research in this

region.

Studies of functional measures of the impact of fire on

microbes in the central hardwoods are more common than

are direct community and density analyses. Vance and

Henderson (1984) in a study of the same Missouri sites later

used by Eivasi and Bryan (1996) found that N mineralization

was reduced by long-term burning, but possibly not with just

a single burn. As there was no change in soil organic C

content due to fire, Vance and Henderson (1984) concluded

that this long-term change was the result primarily of a

change in the quality of the organic C in the soil. In contrast

to what Vance and Henderson (1984) concluded, a

progressive reduction in microbial activity or biomass is also

a possible cause for this long-term reduction in N

mineralization, and this is what Eivasi and Bryan (1996)

reported 12 years later. Eivasi and Bryan (1996)

demonstrated that microbial biomass was reduced relative to

the control by 32% in the annual burn plots and by 21% in

the periodic burn plots. In addition, Eivasi and Bryan (1996)

demonstrated significant, burning-induced reductions in the

activity rates of five key enzymes that serve as indicators of

microbial activity: Acid phosphatase, a- and b-glucosidase,

sulfatase, and urease. Vance and Henderson (1984) also

report long-term reductions in tree growth on the burned

plots, and attribute this to reduced N mineralization, thus

linking the microbial effects to above ground ecological

impacts.

In contrast to the Missouri case, studies done in many

temperate ecosystem types have demonstrated an increase

in N mineralization and N availability after a single or small

number of fires (Boerner 1982). These increases are often

attributed to the alteration of the organic matter by fire in

such a way as to render it more susceptible to microbial

decay, to increased microbial activity, and to altered

microclimate. For example, Boerner et al. (2000b)

demonstrated strong increases in N mineralization in plots
62 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

burned thrice relative to those in those unburned plots. Plots

burned once were intermediate in N mineralization rate.

However, this observed increase in the activity of those fungi

and bacterial involved in N mineralization probably cannot be

generalized to all soil microbial groups. Both Saa et al.

(1993) in pine forests and Boerner et al (2000a) in oakhickory forests report decreased acid phosphatase activity

following a single fire.

Summary and Conclusions

Fires in the central hardwoods are typically low in intensity

and consume primarily the unconsolidated leaf litter. As long

as the fire can move across the open forest floor, soil

temperatures generally do not increase enough to cause

significant heating-induced mortality of organisms dwelling in

the mineral soil. Soils under smoldering piles of woody fuels

may, however, be subject to sterilization. Direct N

volatilization is probably not an important pathway for firerelated nutrient loss due to low fire temperatures. The

microclimate at the forest floor surface is probably affected

significantly and this may produce phenological changes in

root growth and microbial activity. More research is in this

area is warranted. Base cations released from dissolving ash

may or may not increase soil pH and cation availability,

depending on the nutrient status of the soil and the amount

of ash deposited. Nitrogen availability typically increases

after one or a small number of fires but may decrease over

the long term. Abundances of soil animals in the forest floor

are reduced by fire whereas those in the mineral soil are

affected little. Recolonization of the redeveloping post-fire

forest floor is rapid. Microbial abundances in the forest floor

are typically reduced by fire, but rapid recolonization by

these groups is also likely, except under smoldering piles of

woody fuels. The database for the central hardwoods region

is scanty and this area is greatly in need of additional

research attention. Based on this scanty database, overall

impacts of low intensity, dormant season fires in the central

hardwoods is less that the impact on above ground

organisms and ecological processes.

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Department of Agriculture, Forest Service.
64 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

1Research Ecologist, USDA Forest Service, Northeastern

Research Station, 359 Main Road, Delaware, OH 43015

2Ecologist, The Nature Conservancy, Ohio Chapter, 6375

Riverside Drive, Suite 50, Dublin, OH 43017

Current address: USDA Forest Service, Rocky Mountain

Research Station, Fire Sciences Laboratory, P.O. Box 8089,

Missoula, MT 59807

Abstract

Oak-dominated ecosystems throughout the central

hardwood region (CHR) are experiencing a shift in species

composition toward more mesic and/or fire-sensitive species

such as red maple (Acer rubrum), sugar maple (Acer

saccharum), and blackgum (Nyssa sylvatica). Prescribed fire

can enhance oak regeneration but few studies have

examined the effects of fire on other ecosystem components,

including understory vegetation. In 1994, a multi-disciplinary

project was initiated to study the use of prescribed fire as a

restoration tool in mixed oak ecosystems in southern Ohio.

Here, I report on the response of understory vegetation after

3 years of early spring (dormant-season) fires. Each year,

from 1995 to 1998, species’ frequencies were recorded in

1,728 2m2

 quadrats in four 75 to 100 ha study areas. Each

study area included a control, infrequently burned (1X), and

frequently burned (3X) unit. In all, 428 vascular plant species

were identified. Surface fires have altered the understory

environment by greatly reducing the quantity of leaf litter on

the forest floor, but light availability has increased only

slightly as the tree canopy has remained closed. Several

groups of species have responded consistently to fire,

including seed bank increasers, tree seedlings, and spring

monocots. On control plots, few species have increased or

decreased in frequency by more than 5 percent. On 1X plots,

18 species have increased and 5 have decreased; on the 3X

plots, 22 species have increased and 18 have decreased in

frequency. From 1995 to 1998, species richness (nearly all

native species) increased on the burned plots relative to the

control plots. This was particularly true at the quadrat level,

where richness has decreased slightly on the control plots

but increased by 11.4 and 10.4 percent on 1X and 3X plots,

respectively. From our research in Ohio and from other

studies in the CHR, we conclude that dormant-season

prescribed fire may be used to promote oak and pine

regeneration while maintaining or increasing the diversity of

native understory species.

Introduction

Fire has played an important role in shaping the structure

and composition of vegetation in the central hardwoods

region (CHR). Fire use by Native Americans is widely

believed to have created and maintained many of the more

open plant communities in the CHR, including savannas,

barrens, and prairies (Baskin and others 1999, Fralish and

others 1999, Heikens 1999). Fire history studies using

dendrochronology methods support the hypothesis that

frequent fire maintained savanna vegetation in the CHR

(Guyette and Cutter 1991, Batek and others 1999).

Currently, fire is being used in the CHR by land management

agencies including the Nature Conservancy, state heritage

programs, USDA Forest Service, and the National Park

Service to restore and maintain these plant communities,

many of which contain threatened and endangered species

(e.g., Bender, this volume)

It is also hypothesized that frequent fire maintained the

dominance of oak (Abrams 1992, Abrams, this volume) and

pine (Waldrop, this volume) in the presettlement CHR

landscape. The absence of fire has been implicated in the

widespread abundance of shade-tolerant and fire-sensitive

tree species in the regeneration layer of oak forests in the

eastern United States (Lorimer 1984, Loftis and McGee

1993, Lorimer and others 1994). Prescribed fire has been

used to regenerate oaks with varying degrees of success

(Wendel and Smith 1986, Barnes and Van Lear 1998, Elliot

and others 1999). However, shelterwood harvests followed

by high intensity prescribed fires have improved oak

regeneration on productive upland sites in Virginia (Brose

and others 1999, Van Lear and Brose, this volume).

In addition to understanding the effects of fire on tree

regeneration, managers need to understand the effects of

fire on the nontree components within the CHR. In this paper

I examine the effects of fire on the understory vegetation

layer (herbaceous and woody species) in forests in and

around the CHR by presenting data from previous studies

and an ongoing study of prescribed fire in mixed-oak forests

in southern Ohio.

Effects of Fire on Understory Vegetation

in the Central Hardwood Region

The understory vegetation layer in central hardwood forests

is composed of several major life forms: graminoids

(grasses, sedges, and rushes), forbs (nongraminoid

herbaceous plants), shrubs, woody vines, and seedlings of

tree species. Herbaceous species, particularly perennial

forbs, comprise the majority of the species pool in CHR

forests (Hutchinson and others 1999). The cover and

abundance of herbaceous species varies annually due to

climatic conditions (Rogers 1983) and throughout the

growing season as spring ephemerals, summergreen

species, and late-summer species emerge and reach

maximum biomass at different times (Bratton 1976, Goebel

and others 1999).

Understory composition and richness are strongly related to

topographic moisture and fertility gradients (Huebner and

others 1995, Olivero and Hix 1998, Hutchinson and others

Fire and Understory Vegetation: A Large-scale Study in Ohio and a Search

for General Response Patterns in Central Hardwood Forests

Todd F. Hutchinson1 and Steve Sutherland 2
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 65

1999), soil parent material (Howell and Vankat 1981), and

land-use history (Motzkin and others 1996), including fire

history (Swan 1970). Mean species richness of 14.5 species

per m2

 has been reported in mixed mesophytic forests in the

southern Appalachians (Duffy and Meier 1992). In the study

described here, mean richness per 2 m2 ranged from 5.9 to

29.6 and averaged 16.7 (Hutchinson and others 1999).

Due to the limited number of fire studies in the CHR, it is

useful to first examine the long-term effects of fire on

understory vegetation from 2 field studies in southeastern

coastal plain pine forests. After 43 years of fire treatments in

loblolly pine (Pinus taeda) stands on the Santee

Experimental Forest, which included control (no fire), and

both periodic and annual winter and summer burns, the

structure, composition, and diversity of understory vegetation

varied primarily by fire treatment (Lewis and Harshbarger

1976, White and others 1991). There was a gradient from

shrub and hardwood domination on control plots to

herbaceous (forb and graminoid) dominance in the burned

plots. Maximum abundance and diversity of native

herbaceous species occurred in plots burned annually

(White and others 1991). Brockway and Lewis (1997)

reported that longleaf pine (P. palustris) stands burned

frequently over 4 decades had a much more diverse and

productive herbaceous understory than control stands, which

developed a dense shrub layer of gallberry (Ilex glabra) that

reduced diversity.

In a long-term study within the CHR, DeSelm and others

(1973) and DeSelm and Clebsch (1991) documented the

effects of annual and periodic fires on vegetation over a 25-

year period (1965-89) in an oak barrens community in middle

Tennessee by recording vegetation cover annually using the

point-intercept method. The burned plots maintained high

forb and graminoid cover; forb cover averaged 46.5 and 28.3

percent on annually and periodically burned plots,

respectively, and graminoid cover averaged 59.6 and 48.3

percent on annual and periodic burn plots. In sharp contrast,

the control plots rapidly lost herbaceous cover, which had

decreased to less than 1 percent by 1976 as woody plants

invaded and shaded the ground-layer vegetation.

In more closed-canopied oak and pine forests, studies on fire

effects have generally consisted of 1 to several fires. The

response of understory vegetation in and around the CHR is

summarized in Table 1.

Species groups that have increased in frequency,

abundance, or cover following fire include annual forbs such

as fireweed (Erechtites hieracifolia) (Masters 1991, Dolan

1994, Sparks and others 1998), legumes (e.g., Desmodium

spp., Lespedeza spp) (Ducey and others 1996, Sparks and

others 1998), warm-season grasses, including Panicum spp.

and little bluestem (Schizachyrium scoparium) (Dolan 1994,

Sparks and others 1998, Elliot and others 1999), and latesummer forbs such as the asters (Aster spp.), goldenrods

(Solidago spp.), and sunflowers (Helianthus spp.) (Swan

1970, Wilhelm and Masters 1994, Sparks and others 1998).

The most consistent species group was the brambles

(Rubus spp.), which increased via abundant germination in

almost every study (e.g., Swan 1970, McGee and

others1995, Nuzzo and others 1996). For all of these

species, increased light availability following fire likely was

the most important response factor, i.e., it increased seed

germination (Pickett and Baskin 1973, Baskin and Baskin

1988) and/or growth and reproduction (Pitelka and others

1985).

Most species that decreased consistently were tree

seedlings, including red maple (Acer rubrum) and black

cherry (Prunus serotina) (Swan 1970, Cole and others

1990), and shrubs and vines, including Virginia creeper

(Parthenocissus quinquefolia) and greenbriar (Smilax

rotundifolia) (Wilhelm and Masters 1994, Nuzzo and others

1996; Arthur and others 1998). The cover of mountain laurel

(Kalmia latifolia), an evergreen shrub, decreased in several

studies, but resprouted prolifically (McGee and others 1995,

Ducey and others 1996, Elliot and others 1999).

Herbaceous cover generally increased following fire (McGee

and others 1995, Nuzzo and others 1996) while the cover

(Nuzzo and others 1996; Elliot and others 1999) and density

(Schwartz and Heim 1996) of woody vegetation decreased

(Table 1). However, due to resprouting, understory woody

cover also has exceeded pre-burn levels within several years

(McGee and others 1995, Arthur and others 1998). When

mixed oak-pine stands in eastern Oklahoma were thinned

and burned, herbaceous biomass increased significantly

(Masters and others 1993).

In nearly all of the fire studies in the CHR, species richness

or diversity increased following fire, though the magnitude

and duration of the responses were highly variable (Table 1).

In a sand forest in Illinois, species richness increased from

6.2-7.0/ m2

 to 9.8-10.8/ m2 following the first of 3 fires, due to

an increase in herbaceous species, particularly annuals

(Nuzzo and others 1996). However, the strong effects were

short-lived as richness subsequently declined after the first

fire-free year. In western Arkansas, species richness

increased significantly following a single dormant-season fire

in restored shortleaf pine (Pinus echinata) forests (Sparks

and others 1998). The stands gained 14-16 species and

more than 3 species were gained per m2

 (Sparks and others

1998). More than 8 years following 2 fires in a transition oaknorthern hardwood forest, forb richness increased

significantly over pre-burn levels in 2 stands, 26 to 44 and 23

to 27 species/4 m2

, respectively (McGee and others 1995).

Other studies reported increased species richness following

fire (Cole and others 1990, Wilhelm and Masters 1994, Taft

and others 1995, Arthur and others 1998), though Dolan

(1994) found no change in richness and Schwartz and Heim

(1996) reported a decrease (Table 1).

Fire intensity and season are other important factors that

influence the response of vegetation. Ducey and others

(1996) and Elliot and others (1999) reported that species

richness increased more following high-intensity fires than in

areas that burned at lower intensities. In Arkansas, dormantseason burns in March and April increased the abundance of

several Panicum species but a September fire reduced

Panicum abundance (Sparks and others 1998). After an
66 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 Table 1.—Summary of fire studies examining the response of understory forest vegetation in and around the ce

Overstory Duration Fire Herbaceous dominant Location (pre-burn data) Treatments response Post oak OK 5 years (yes) 1 or 3 fires richness + Shortleaf pine Winter Shortleaf pine AR 2 years (yes) 1 or 2 fires richness + 

Mar-Apr; Sep density + / - Mixed: oak, pine, NC 2 years (yes) 1 fire richness + 

maple, hickory Apr cover + Scarlet oak KY 4 years (no) 1 or 2 fires richness + Chestnut oak Mar cover + Post oak IN 7 years (yes) 1 fire Richness: 

Apr No effect Mixed oak NY 1 year (no) 1 wildfire # increasers > Apr-May # decreasers Red oak, white CT 1 year (no) 1 fire richness + 

oak, white pine Apr density + Red oak, NY 12 years (yes) 1 or 2 fires richness + white oak Apr height + Black oak IL 5 years (yes) 3 fires (dormant) richness + 

autumn and spring cover + White oak, red IL 4 years (yes) 1 fire richness - 

oak, black walnut Mar or May (May burn only) Mesic woodland IL 5 years (no) 4 fires richness + 

Spring
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 67

initial depression in species richness following both March

(dormant season) and May (growing season) fires in Illinois,

richness soon recovered following the March but not the May

burns (Schwartz and Heim 1996).

Fire has not facilitated the invasion of exotic species in CHR,

possibly because most of the areas that have been burned

and studied quantitatively have been relatively xeric and

undisturbed, and thus not highly invasible (Rejmanek 1989).

In more mesic and disturbed forests where garlic mustard

(Alliaria petiolata) has invaded, fire moderately reduced both

its cover and abundance (Nuzzo and others 1996, Schwartz

and Heim 1996).

Effects of Fire on Understory Vegetation

in Ohio Oak Forests

Witness tree data from land surveys indicates that the

unglaciated Allegheny Plateau region of southeastern Ohio

was composed primarily of mixed oak forests just prior to

European settlement (ca. 1800) (Gordon 1969). In the

southern portion of this region, many of the oak forests were

harvested repeatedly from 1850 to 1900 to supply charcoal

for the numerous iron furnaces in the region (Stout 1933). In

secondary forests dating to the end of the iron-producing

period, oaks, primarily white (Quercus alba), chestnut (Q.

prinus) and black (Q. velutina), continue to dominate the

overstory but the midstory and understory layers are

dominated by shade-tolerant species such as red maple

(Acer rubrum), sugar maple (Acer saccharum), blackgum

(Nyssa sylvatica), and beech (Fagus grandifolia) (Figure 1).

Similar trends have been reported for numerous oak forests

in the eastern United States (Abrams 1992). On the basis of

these forest trends, a multidisciplinary study was established

to examine prescribed fire as a tool to restore structure,

composition, and function to oak forests in southern Ohio.

Methods

Study Areas and Experimental Design

In 1994, four study areas (75 to 100 ha each) were

established, two in Vinton County (Arch Rock, 39o

12' N,

82o

23' W; Watch Rock, 39o11" N, 82o22' W), and two in

Lawrence County (Young’s Branch (38o43' N, 82o

31' W;

Bluegrass Ridge (38o

36' N, 82o31' W). The sites were

dissected, underlain by sandstones and siltstones, and had

moderately acidic siltloam alfisols with low water-holding

capacity. The study areas have been relatively undisturbed

since clearcutting 80 to 120 years ago, and were dominated

by oak. Average tree basal area was 24 to 28 m2

/ha

(Hutchinson and others 1999). Three fire treatment units

were established within each area. Control units were not

burned, infrequent units were burned once (1X) in MarchApril 1996, and frequent units were burned three times (3X)

in March-April, 1996, 1997, and 1998. The surface fires

generally were low intensity, with flame lengths less than 50

cm and temperatures averaging 89.6o

C (193.2oF) at a height

of 25 cm over the 3-year period (Hutchinson, unpublished

data). In the uplands of each unit, nine 50 x 25m (0.125 ha)

vegetation plots (N = 108 total plots) were stratified by a

GIS-derived integrated moisture index based on a suite of

landscape features (Iverson and others 1997).

Field Data Collection

The frequency of understory species was recorded in four 2

x 1 m quadrats along each of four transects spaced 5 m

apart within a 25 x 25m portion of each plot (16 quadrats per

plot). Each year, three quadrats per transect were located

randomly and one quadrat was located permanently at the

midpoint of each transect. Here, data from random and

permanent quadrats were combined. Vegetation was

sampled twice each year (May and September) from 1995

(pre-burn) to 1998. Herbaceous plants were recorded on a

presence/absence basis in each quadrat to obtain frequency.

For each species, the maximum frequency was recorded for

the year. For understory individuals of tree species, we

recorded the presence of each species in each of four size

classes: < 30 cm height, 30 cm height to 2.99 cm diameter at

breast height (dbh), 3.0 to 9.9 cm dbh, and > 10 cm dbh.

Figure 1.—Forest composition trends in southern Ohio. Witness tree

data are from townships in and adjacent to the study areas. Data for

the Vinton County areas are from Beatley (1959). Current overstory

(trees > 10 cm dbh) and sapling (trees 1.37 m height to 9.9 cm dbh)

data were collected from 108 vegetation plots The primary “other”

species for the saplings are blackgum and beech.
68 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Results and Discussion

Following the first fires, 37 percent of the small saplings

(1.37 m height - 2.9 cm dbh), 9 percent of the medium

saplings (3.0 - 5.9), and 4 percent of the large saplings (6.0 -

9.9) had been topkilled though most stems resprouted

(Sutherland and others 1997). After three fires, nearly 95

percent of the small saplings had been topkilled (Sutherland

and Hutchinson, unpublished). However, few trees greater

than 10 cm dbh have been topkilled (D. Yaussy, USDA

Forest Service, unpublished) and open sky remained below

6 percent after two fires on the 3X plots (Robison and

McCarthy 1999). Leaf litter consumption ranged from 28.5 to

80.4 percent during the first fires in 1996 (Boerner and

others, in press). Litter mass has continued to decrease on

frequently burned plots, while increasing to pre-burn levels

by 1998 on the 1X plots (Hutchinson, unpublished).

Over the 4-year period, we recorded 428 vascular plant

species, composed of 254 perennial forbs, 59 graminoids,

45 trees, 30 shrubs, 24 annual forbs, 11 woody vines, and 5

monocarpic perennial forbs. While the frequency of most

species did not change significantly following prescribed fire,

the following groups exhibited distinctive patterns of change.

Seed Bank Increasers

Several species increased in frequency following abundant

germination from the seed bank. Fireweed (Erechtites

hieracifolia) and yellow-poplar (Liriodendron tulipifera) had

the greatest increases in frequency on the burned plots

(Figure 2). Prior to burning, fireweed had a mean frequency

of less than 10 percent per plot; following the first fires in

1996, it increased to nearly 80 percent frequency

Presumably, most of the germination was from the buried

seed pool as fires would have consumed the previous

autumn’s small, wind-dispersed seeds. On the 1X plots, the

frequency of fireweed returned to pre-burn levels following 2

years without fire; frequency remained high (> 55 percent)

on 3X plots. Like many annuals, fireweed probably requires

light for germination (Pickett and Baskin 1973, Baskin and

Baskin 1988). Disturbances that remove litter and expose

mineral soil are required to maintain fireweed at high

frequencies in standing vegetation. Yellow-poplar, which

produces seeds that are viable in the seed bank for 4 to 7

years (Burns and Honkala 1990), increased in seedling

frequency from less than 25 percent prior to burning to

nearly 80 percent on burned plots following the first fires. In

contrast to fireweed, the frequency of yellow-poplar

seedlings then decreased to approximately 50 percent on

the 1X and 3X plots from 1996 to 1998. On the 1X plots, the

decrease likely was caused by seedling mortality due to low

light levels (Burns and Honkala 1990). On the 3X plots fire

likely killed most of the seedlings that had germinated,

though new germination maintained relatively high

frequencies of seedlings.

Other species in the pre-burn standing vegetation that

increased in frequency by more than 10 percent on the

burned plots included Virginia copperleaf (Acalypha

virginica), sedges (Carex spp.), white snakeroot (Eupatorium

rugosum), sweet-scented bedstraw (Galium triflorum), panic

grasses (Panicum boscii, P. latifolium), brambles (Rubus

spp.), sassafras (Sassafras albidum), violets (Viola spp.),

and wild grapevine (Vitis spp.) Smooth sumac (Rhus glabra),

which was absent from the pre-burn standing vegetation,

increased in frequency to 11.3 percent on the 3X plots by

1998. Several species not recorded in the pre-burn standing

vegetation germinated following fire, but remained at

frequencies under 10 percent. These species included

sensitive plant (Chamaecrista nictitans), horse-weed

(Conyza canadensis), late eupatorium (Eupatorium

serotinum), upland boneset (E. sessilifolium), cudweed

(Gnaphalium obtusifolium), beggar’s-lice (Hackelia

virginiana), indian-tobacco (Lobelia inflata), pokeweed

(Phytolacca americana), dwarf sumac (Rhus copallina), firepink (Silene virginica), and Sphenopholis nitida, a grass. Of

several exotic species that appeared after the fires, only

black bindweed (Polygonum convolvulus) was found in more

than 1 percent of quadrats within burned treatments (2.4

percent in 1998).

Figure 2.—Frequency data for fireweed and yellow-poplar

seedlings. Mean frequencies were calculated as the mean number of

quadrats per plot in which the species occurred for 36 plots per year

in each of three fire treatments (fires are indicated with an F).
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 69

Tree Seedlings

The effects of fire on tree seedlings varied by species, as

shown by the contrasting responses of red maple and white

oak (Figure 3). From 1995 to 1998, the frequency of red

maple seedlings increased from 63.0 to 70.8 percent on

control plots, but decreased from 67.4 to 43.1 percent on the

3X plots. On the 1X plots, the frequency of red maple

seedlings decreased from 58.4 to 44.6 percent following the

fire but increased to 63.4 percent in 1998 after 2 years

without fire. Although larger red maple seedlings sprout

following topkill, three fires reduced frequencies

substantially. By contrast, seedlings of white oak, which was

the most abundant overstory species and the most frequent

oak species in the seedling layer, exhibited a neutral

response (< 5 percent frequency shift) to fire (Figure 3).

White oak seedlings increased in frequency in all treatments

following a mast year in 1996. In 1998, frequencies were

nearly identical across treatments (28.3 to 30.2 percent).

Other fire studies have shown similar responses of red

maple and oaks (e.g., Kruger and Reich 1997).

Other tree species that decreased in seedling frequency by

more than 5 percent from 1995 to 1998 on the 3X plots were

flowering dogwood (Cornus florida, 17.0 percent), American

ash (Fraxinus americana, 15.8), sugar maple (Acer

saccharum, 10.4), slippery elm (Ulmus rubra, 8.2), hophornbeam (Ostrya virginiana, 7.5), and black cherry (Prunus

serotina 6.9). In addition to white oak, common seedlings

that showed a neutral response to fire on the burned plots

included scarlet oak (Quercus coccinea), chestnut oak (Q.

prinus), red oak (Q. rubra), black oak (Q. velutina), pignut

hickory (Carya glabra), bitternut hickory (C. cordiformis),

redbud (Cercis canadensis), hawthorn (Crataegus spp.), and

blackgum (Nyssa sylvatica).

Spring Monocots

Several monocots that emerge early in the spring decreased

in frequency on the burned plots, particularly in years when

the fires were conducted later in the spring. Both false

Solomon’s seal (Smilacina racemosa) and large-flowered

trillium (Trillium grandiflorum) decreased in frequency by

Figure 3.—Frequency data for red maple and white oak seedlings.

Mean frequencies were calculated as the mean number of quadrats

per plot in which the species occurred for 36 plots per year in each

of three fire treatments (fires are indicated with an F).

Figure 4.—Frequency data for false solomon’s seal and largeflowered trillium. Mean frequencies were calculated as the mean

number of quadrats per plot in which the species occurred for 36 plots

per year in each of three fire treatments (fires are indicated with an F).
70 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Figure 5.—Species that increased or decreased in mean frequency by

more than 5 percent from 1995 to 1998 in three fire treatments.

12.5 to 16.5 percent following the first fires (Figure 4). The

1996 fires were conducted later (through April 22) than in

subsequent years due to a wet spring. However, in both fire

treatments, these species returned to pre-burn frequencies

in 1997 following another fire on the 3X plots and no fire on

the 1X plots. The 1997 burns were completed by April 3,

suggesting that dormant-season fires have a negative impact

on these species only if conducted when they begin to

emerge. Damage to the leaf tips of several monocot species

has been observed frequently in the study areas (personal

observation). From 1997 to 1998 the frequency of false

solomon’s seal and trillium decreased by 6.6 and 5.4

percent, respectively, suggesting that the fires occurred at a

phenological stage intermediate to 1996 and 1997 (Figure 4).

Solomon’s seal (Polygonatum biflorum) and bellwort

(Uvularia perfoliata) also decreased in frequency on burned

plots.

Summary of Increasers and Decreasers

To examine the effects of fire on the entire species pool, we

classified species as increasers or decreasers if their

frequency shifted by more than 5 percent from 1995 (preburn) to 1998. On the controls, few species shifted in

frequency by more than 5 percent (Figure 5). On the 1X

plots, there were 18 increasers, of which 11 were graminoids

and perennial forbs, and 5 decreasers. On the 3X plots,

there were 22 increasers but also 18 decreasers, most of

which were woody species (11), including 8 tree species.

These results indicate that the composition of the 3X plots

has changed more than the1X plots, primarily due to the

reduced frequencies of several tree species. Despite the

greater change on the 3X plots, most species have not

shifted more than 5 percent following dormant-season

prescribed fires. This suggests that perennial herbaceous

species have not been damaged directly by fire, and have
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 71

emerged in numbers similar to those on the controls. Also,

most woody species that were topkilled by fire have

resprouted. Although there have been several seed bank

increasers, most species have not exhibited abundant

germination following fires.

Species Richness

From 1995 to 1998, species richness per quadrat increased

by 11.4 percent on 1X plots and 10.4 percent on 3X plots.

Over the same period, richness decreased by less than 1

percent on the controls (Figure 6a). Species richness per

plot (based on 16 quadrats) also increased more on the

burned plots relative to the controls, but only by several

species; richness increased by 2.4 percent on control plots,

3.4 percent on 1X plots and 5.0 percent on 3X plots (Figure

6b). When richness was summed for the nine plots in each

unit, richness increased in all treatments from 1995-98, but

to a greater degree on the burned plots. Richness per unit

increased by 3.4 percent on control plots, 5.6 percent on 3X

plots, and 8.9 percent on 1X plots. (Figure 6c).

The increases in species richness on the burned plots are

subtle, but our results indicate that fire has not depressed

richness in these oak forests over the initial 4-year period of

this study. Our results also underscore the need for pretreatment data when conducting fire studies. Our control

units were less species rich prior to treatment at all three

scales of observation. Had we examined only post-burn

data, the presumed effects of fire on increased species

richness would have been exaggerated.

Conclusions

Because trees dominate the structure of CHR forests and

have high economic value, most research and discussion on

fire effects in this region has focused on tree regeneration.

However, plant diversity in the CHR region is concentrated in

the understory layer, particularly among herbaceous species.

It is these plants that also provide habitat and food for

diverse assemblages of fauna. There is growing evidence

that plant diversity directly affects the productivity and

sustainability of ecosystems (Tilman and others 1996). From

studies in the CHR it is clear that dormant season fire has no

negative impact on plant diversity. In fact, most studies show

small to moderate increases in diversity as fire creates an

improved regeneration niche for some species (Table 1).

Our Ohio study showed that prescribed fire has both direct

and indirect effects on undestory vegetation. Fire directly

affects species composition by topkilling most of the smaller

woody stems. The relative frequencies of woody species

then shift due to variation in resprouting ability; we reported a

decrease in the relative frequencies of maple, dogwood, ash,

and elm seedlings to those of the oaks and hickories. Fire

also directly damaged several herbaceous species (spring

monocots), causing reduced frequencies.

Indirectly, fire caused a shift in species composition by

greatly reducing leaf litter and exposing mineral soil, which

facilitated the abundant germination of several species from

the seed bank and the addition of species to the standing

vegetation. Other indirect fire effects that may affect species

composition include a subtle increase in light availability

throughout the growing season, an increase in soil nutrients

and pH from leaf litter ash (see Boerner, this volume), and a

decrease in soil moisture through increased evaporation.

It is clear that fire suppression since the 1940’s has resulted

in significant changes in CHR forests. There is a need for

long-term research on the effects of fire on the understory

vegetation in these forests and for additional population-level

studies of fire response by individual species. We also need

to understand how the effects of fire and fire suppression on

understory vegetation are linked to physical and biological

patterns and processes (e.g., light availability and nutrient

cycling) and to the population dynamics and behavior of

animals, including herbivores, pollinators, and microbial

communities.

Figure 6.—Mean species richness values for the three fire

treatments at three spatial scales of observation (fires are

indicated with an F).
72 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Acknowledgments

We thank the many individuals and agencies whose

assistance was invaluable in conducting this project. Marilyn

Ortt provided the botanical expertise as well as 4 years of

field work, Elaine Kennedy Sutherland provided project

leadership, and Marty Jones, Ralph Boerner, and Louis

Iverson reviewed an earlier draft of this manuscript. The

following participated in either the collection of field data and/

or assisted with data entry: Heather Bentsen, Rosemarie

Boyle, James Buratti, Ginger Burnhold, Carolee Franklin,

Rick Gardner, John Gire, Jennifer Gray, Meryl Hattenbach,

Jennifer Hillmer, Erin Hines, David Hosack, Cindy Huebner,

Louis Iverson, Julie Lapp, John Leach, Bob Long, Dave

Minney, Anantha Prasad, Ava Rawn, Mary Ann Tate, and Jim

Yerian. Mary Boda assisted with the graphical presentation

of the data for the conference. We also thank Wayne

Lashbrook, Steve Mathey, and Walt Smith, Mead Paper

Corp., and Ralph Miller, Wayne National Forest, for

supplying the study sites and logistical support. For

conducting the prescribed fires, we thank personnel with the

Wayne National Forest, particularly Art Martin, Terry Krasko,

Tom Eaches, and Nelson Kruger who served as burn

bosses. The following agencies and institutions also

contributed personnel to conduct the fires: The Ohio Chapter

of the Nature Conservancy, The Ohio Department of Natural

Resources, Divisions of Forestry and Natural Areas and

Preserves, Hocking Technical College, The Ohio State

University, and Ohio University. Funding was provided by the

Washington Office of the USDA Forest Service in a grant to

Elaine Kennedy Sutherland for ecosystem management

research.

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Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 75

Abstract

Fire is often viewed as destructive of wildlife and wildlife

habitat, because it can cause dramatic change. The

perception of catastrophic fire that burns up forests and kills

wildlife has been fostered to enlist public support for

suppressing fires. A message that is much more difficult to

convey is that when fire is removed from the landscape

important wildlife habitat can be slowly and subtly eliminated

from ecosystems that depend on the frequent occurrence of

fire.

Many fire-dependent communities—shortleaf pine forests,

barrens, glades, prairies, balds, and oak savannas—are

embedded in the extensive and diverse central hardwood

landscape. Approximately 232 species of birds (119

breeding, 44 wintering, and 69 year-round residents) reside

in this landscape. Some of the rarer bird species in the

region are closely tied to fire-dependent habitats. The status

of the Red-cockaded Woodpecker (Picoides borealis) is

tenuous in pine forests of Kentucky and Oklahoma where fire

has been suppressed for much of the century. Efforts to

reintroduce fire to restore the open-structured, pinedominated forests are hopeful signs for the species.

Grassland birds are one of the most rapidly declining groups

of species in North America. Habitat conversion has

eliminated many of the grassland communities within the

Central Hardwoods region, but plant succession in the

absence of fire has also degraded habitat suitability for many

grassland birds. Numerous preserves are being established

to retain the rare biota of these grasslands. Many oakdominated forests and oak woodland- savanna mosaics are

slowly shifting to greater proportions of trees more

associated with mesic conditions, such as maples (Acer sp.),

and canopy closure. These shifts in structure and

composition bring attendant changes in the avian

community. Fire—with proper consideration of the potential

effects on small populations of birds and invertebrates and

the tradeoffs in bird species that will be favored—is an

essential tool for habitat maintenance for some of the rare

and declining birds in the Central Hardwoods region.

Introduction

A diverse array of vegetation communities that have

experienced frequent fire for millennia are scattered

throughout the extensive Central Hardwoods forest region

(Hicks 1998; Braun 1950; Braun 1950; Hicks 1998; Anderson

and others 1999). Some of these communities were

fragmented and highly dispersed (e.g., balds) and some

were extensive (e.g., the Kentucky barrens). The structure

and plant composition of some of these communities was

largely shaped edaphic conditions (Quarterman and others

1993), but fire played an important role to arrest succession

and control brush and saplings as soils developed (Deselm

and Murdock 1993). Lightning-started fires played a role in

creating some of these habitats, but Native Americans

undoubtedly used fire frequently and pervasively to create

the open habitats found by early European colonists

(Bonnicksen and others 1999; Askins 1999). Timber harvest,

fire suppression, and plant succession have dramatically

changed the composition of some of these plant

communities, and, in the process, the avian comunities.

Excellent descriptions of some of the grassland preserves in

the Central Hardwoods region (e.g., Tennessee) can be

found at: http://www.tnc.org/infield/State/Tennessee/

The purpose of this paper is to provide a brief overview of

the avifauna of the region, including species that have

become extinct or extirpated, to explore the bird species

composition of three general vegetation communities that

were shaped by frequent fires—shortleaf pine (Pinus

echinata) forests, grassland/shrubland communities (balds,

barrens, and glades), and open oak woodlands and

savannas—and to discuss the effects on birds of use of

prescribed fire in the management and restoration of these

communities.

The Central Hardwoods Avifauna

Hicks (1998) uses four of Braun’s (1950) forest types to

define the Central Hardwoods region: the mixed mesophytic,

western mesophytic, oak-hickory, and oak-chestnut forests.

This large region extends over all or part of 35 states (Figure

1). I obtained a list of the bird species within this region by

comparing distribution maps of bird species (Dunn 1987) to

the map of the Central Hardwoods. Bird species whose

winter or breeding season distributions overlapped with the

outline of the Central Hardwoods region were included;

migrant and transient birds were not.

Approximately 232 bird species winter or breed within the

Central Hardwoods region (Appendix); all scientific names

follow the American Ornithologists’ Union (1998). Sixty-nine

(30%) of these species occur year-round, 44 (19%) occur in

the winter, and 119 (51%) occur in the breeding season only.

Four bird species that were found in the Central Hardwoods

historically, Passenger Pigeon (Ectopistes migratorius),

Carolina Parakeet (Conuropsis carolinensis), Ivory-billed

Woodpecker (Campephilus principalis), and Bachman’s

Warbler (Vermivora bachmanii), have become extinct. The

Passenger Pigeon and Bachman’s Warbler may have been

affected by changes in fire regimes in the Central

Hardwoods region, although it seems likely that outright

habitat loss was more important in their decline than habitat

degradation through fire exclusion. For example burning in

forests by Native Americans kept forest structure open and

Fire and Birds in the Central Hardwood Landscape

R. Todd Engstrom1

1Tall Timbers Research Station, 13093 Henry Beadel Road,

Tallahassee, FL 32312-0918
76 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

parklike, which may have favored mast producing species

and facilitated foraging by Passenger Pigeons on the

ground. Bucher (1992) has hypothesized that destruction of

forest breeding habitat and the lack of social facilitation in

foraging were the major factors that contributed to the

extinction of the species. Bachman’s Warbler may have

narrowly specialized on canebrakes (Arundinaria spp.) in

bottomland hardwoods for breeding habitat (Remsen 1986).

Fire played a critical role in maintenance of these large

stands of cane (Hughes 1966).

Several bird species have been nearly extirpated from the

Central Hardwoods landscape. The distribution within the

region of Greater Prairie Chicken, Red-cockaded

Woodpecker, Bewick’s Wren, and Brown-headed Nuthatch

have all receded within the last 200 years. The Greater

Prairie Chicken formerly occurred in Ohio, Kentucky,

Arkansas, and Missouri, and the eastern seaboard from

Massachusetts and Virginia, but disappeared from hunting

pressure and habitat destruction (Schroeder and Robb

1993). Lumsden (1966) suggested that the Greater Prairie

Chicken may have colonized the eastern U.S. in response to

grassland openings created when Native Americans burned

woodlands.

Causes for the decline of the Bewick’s Wren in the Central

Hardwoods is far from clear. The most common explanation

is that the Bewick’s Wren expanded eastward in the early

19th century in response to agricultural expansion, but was

then out-competed by the House Wren as it expanded its

range (Kennedy and White 1997). Although the Bewick’s

Wren was not found in the region prior to 1821, C. Hunter

(pers. comm.) has suggested that it might have been present

in the oak-savanna mosaic of the Central Hardwoods. Loss

of this habitat to agriculture and modification of the fire

regime may have degraded the best habitat for the Bewick’s

Wren.

Pine Woodland Specialists

in the Central Hardwoods Region

Some of the rarest and most threatened bird species of the

Central Hardwoods landscape are associated with pine

woodlands along the southern edge of the region. The Redcockaded Woodpecker, a federally endangered species

since 1970, is highly associated with open pinelands in the

southeastern United States (Jackson 1994). Historically, the

distribution of the woodpecker within the Central Hardwoods

was primarily the Ouachita Highlands of Arkansas, Missouri,

and Oklahoma, and the Cumberland Plateau of Kentucky

and Tennessee (fig. 1). This species was typically associated

with stands of shortleaf pine (Pinus echinata) on dry slopes

and ridges. Interestingly, the distribution of Red-cockaded

Figure 1.—Central Hardwoods region from Braun (1950).
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 77

Woodpecker clusters in the Daniel Boone National Forest

has striking overlap with the distribution of Native American

archeological sites (L. Perry, pers. comm.), which suggests

that Native American use of fire may have been important in

maintaining habitat for the woodpecker.

The Red-cockaded Woodpecker was apparently extirpated

from Missouri by 1946 (Eddleman and Clawson 1987) and

from Tennessee by 1994 (Nicholson 1997). Only one

individual native to Kentucky remains, although 16

individuals have been translocated to Kentucky from larger

populations to support the recovery effort there (L. Perry,

pers. comm.). In each of these states the loss or decline of

the red-cockaded woodpecker seemed to follow a similar

pattern: elimination of old-growth shortleaf pine and drastic

reduction of the fire frequency put extreme pressure on

populations that were naturally small and fragmented by

topography.

The largest remaining population of Red-cockaded

Woodpeckers in the Central Hardwoods region is found in

the 5701-ha McCurtain County Wilderness Area in

southeastern Oklahoma, which is dominated by old-growth

shortleaf pine-hardwood forest on steep ridges. Historical

accounts characterize the vegetation of the Ouachita

Mountains of Oklahoma “...as widely spaced trees with low

basal area and stem density, open grassy understories,

sparse midstories, and indicate that fire shaped forest

conditions (e.g., pine-bluestem communities)” (Masters and

others 1995). Substantial evidence indicates that Native

Americans (the Caddo Indians before 1751 and the Choctaw

in the mid-nineteenth century) used fire frequently in the

area. Fire was also used frequently by early European

settlers as indicated by analysis of fire scars, up until the era

of modern fire suppression in 1957 (Masters and others

1995). Mean fire interval of the Wilderness Area during the

presettlement period was between 3.5 and 5.6 years; this

extended to over 500 years in the era of modern fire

suppression.

Fire suppression and attendant extensive growth of

hardwoods within the McCurtain County Wilderness Area is

believed to have been partially responsible for the decline of

the Red-cockaded Woodpecker (Kelly 1991; Masters and

others 1995; Masters and others 1995). Wood (1977)

located 29 groups (89-92 individuals) from 1974-77; Kelly

(1991) found 15 active clusters (31 individuals); and Masters

and others (1995) reported 9 active clusters (22 individuals):

an average loss of 1.4 clusters per year. The Oklahoma

Department of Wildlife Conservation is actively working to

restore woodpecker habitat through hardwood control and

reintroduction of fire.

Efforts to restore Red-cockaded Woodpecker habitat include

removal of hardwoods that have grown in response to pine

harvest and fire suppression and reintroduction of fire. In the

Central Hardwoods region Red-cockaded Woodpecker

habitat restoration has resulted in predictable effects on

other species in the avian community. (Wilson and others

1995) reported increases in numbers of 10 pine-grassland

bird species (Eastern Wood-Pewee, Northern Bobwhite,

Red-headed Woodpecker, Prairie Warbler, Indigo Bunting,

Bachman’s Sparrow) in response to management for Redcockaded Woodpeckers in Arkansas. (Hines 1999) found

similar response of the avian community in Kentucky. In both

studies some “interior” species (Black-and-white Warbler,

Ovenbird) were found more frequently in control plots than in

treatments.

Avian Communities of Savannas, Barrens,

and Glades in the Central Hardwoods Region

Savannas, barrens, and glades were important components

of the Central Hardwoods landscape before European

settlement and even before human colonization. Pine

barrens in New Jersey and New York (Gibson and others

1999), serpentine barrens in Pennsylvania (Tyndall and Hull

1999), rock outcrop communities (Quarterman and others

1993), cedar glades (Baskin and Baskin 1999), and the Big

Barrens region of Kentucky and Tennessee (Baskin and

others 1999) are some of the vegetation types that provided

open, grassland/shrubland/savanna habitats for birds in the

Central Hardwoods region. Some of these plant communities

were extensive--the Big Barrens was approximately 2 million

acres at the time of European settlement–but many were

relatively small and isolated (Quarterman and others 1993)

historical and prehistorical evidence suggests that

grasslands in the region have long provided important

habitat for many species of birds (Askins 1999).

Many of these communities share common features that

inhibit tree growth: harsh edaphic conditions (i.e., shallow

soils over bedrock that dry quickly) and the frequent

occurrence of fire (Anderson and others 1999). Use of fire by

Native Americans played an essential role in retarding

succession and maintaining these communities with few or

no shrubs or trees (Bonnicksen and others 1999; Askins

1999).

Populations of birds, as measured by the North American

Breeding Bird Survey (BBS), that inhabit grassland,

shrubland, and savanna communities are showing some of

the most dramatic declines of any functional group of bird

species in the eastern United States (Peterjohn and Sauer

1999). To better focus on bird population trends in the

Central Hardwoods region, I selected the 10 physiographic

regions within the Central Hardwoods region that are used

by the BBS for trend estimates (table 1). I counted the

number of physiographic regions in which each species

showed statistically significant positive and negative trends

over the period 1965 to 1995 (DeGraaf and Rappole 1995;

Appendix). For example the Field Sparrow has had

significant negative trends over a 30-year period in 9 of the

10 BBS physiographic regions; the American Robin has

significantly increased in 7 of 10 regions (Appendix).

Relatively few studies have been published of the avian

communities of the savannas, barrens, and glades of the

Central Hardwoods region. One study of a cedar glade

(Schultz 1930; Quarterman et. al 1993) and a Breeding Bird

Census in a serpentine grassland in Pennsylvania

(Greenewalt and others 1961) indicate that many of the birds
78 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

of these vegetation communities are showing significant

regional declines (table 2).

Some of the best remaining examples of grasslands in the

Central Hardwoods region have some form of conservation

protection. Management objectives of these preserves

frequently include maintenance of small, relatively isolated

populations of rare plants, invertebrates, and birds. Burning

in grassland preserves such as these requires special

attention (Engstrom 1997). Consider the following general

suggestions for managing grasslands. (1) Mix up

management applications (e.g., season of fire) from year-toyear to provide variation in disturbance. No single

management prescription will be best for all species. (2)

Consider vegetation litter recovery and accumulation when

establishing fire frequency. This is important for some bird

species, such as Henslow’s Sparrow, which prefers relatively

thick grasses for nesting. (3) Consider the size of the

management area in relation to the needs of the species of

interest. Subdividing large tracts into smaller units and

applying different management practices on different units

would provide habitats in a variety of stages of recovery from

disturbance.

Birds of Oak Savannas

Fire is critical for determining the structure and composition

of vegetation of sand savannas (Will-Wolf and Stearns

1999), deep soil savannas (Anderson and Bowles 1999),

and open oak woodlands (Fralish and others 1999) in the

Central Hardwoods region. In the absence of fire, shadetolerant tree species such as sugar maple (Acer saccharum),

gradually replace oaks (Abrams 1992) close the canopy, and

create more mesic conditions. Changes in tree species

composition and forest structure strongly affect the avian

community. Recently, (Brawn 1998) and (Artman and

Downhower 2000) have experimentally studied the effects of

fire on bird communities in oak woodlands in Illinois and

Ohio respectively.

As in the cases of restoration of shortleaf pine woodlands the

use of fire to manage oak woodlands and savannas favors

some bird species over others (table 3). Note that many of

the bird species that increase significantly after application

are those that are declining within the region.

Conclusions

Many of the structurally open habitats in the Central

Hardwoods region probably originated because edaphic

Table 1.—Comparison of physiographic regions used by Braun (1950) and the Breeding

Bird Survey (Robbins et al. 1986).

Braun (1950) Breeding Bird Survey

1. Mixed Mesophytic Forest 21. Cumberland Plateau

a. Cumberland Mtns. 22. Ohio Hills

b. Alleghany Mtns.

c. Cumberland and Allelghany Plateaus

2. Western Mesophytic Forest

a. Bluegrass 14. Highland Rim

b. Nashville basin 15. Lexington Plain

c. Illinoian glaciation

d. Mississippian Plateau

e. Mississippi Embayment

3. Oak-hickory Forest

 Southern Division 19. Ozark-Ouachita Plateau

a. Interior Highlands 31. Till Plains

b. Forest-Prairie Transition

 Northern Division

c. Mississippi Valley

d. Prairie Peninsula

4. Oak-Chestnut Forest

a. Southern Apalachians 10. Northern Piedmont

b. Northern Blue Ridge 12. Southern New England

c. Ridge and Valley 13. Ridge and Valley

d. Piedmont 23. Blue Ridge Mountains

e. Glaciated
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 79

Table 2.—Bird species found in two grassland communities in the Central Hardwoods region.

Underlined species are showing significant declines in at least three physiographic regions

used by the Breeding Bird Survey (See Appendix).

Cedar glade in central Tennessee Serpentine Barrens in Pennsylvania

(Schultz 1930) (Greenewalt et al. 1961)

Northern Cardinal Rufous-sided Towhee

Indigo Bunting Field Sparrow

Field Sparrow Gray Catbird

Chipping Sparrow Brown Thrasher

Rufous-sided Towhee

White-eyed Vireo

Red-eyed Vireo

Carolina Chickadee

Common Yellowthroat

Carolina Wren

Blue Jay

Yellow-breasted Chat

American Crow

Tufted Titmouse

Wood Thrush

Summer Tanager

Blue-gray Gnatcatcher

Northern Mockingbird

Northern Flicker

Brown Thrasher

Gray Catbird

Northern Bobwhite

Mourning Dove

Yellow-billed Cuckoo

Downy Woodpecker

Eastern Bluebird

Orchard Oriole

Table 3—Bird species that showed significant responses to use of prescribed fire in

oak woodlands.

Artman and Downhower

Brawn (1998) (this conference)

Northern Bobwhite Positive

Mourning Dove Positive

Red-headed Woodpecker Positive

Great Crested Flycatcher Positive

Eastern Wood-Pewee Positive

American Robin Positive

Wood Thrush Negative

Veery Negative

Brown Thrasher Positive

Ovenbird Negative Negative

Hooded Warbler Negative

Worm-eating Warbler Negative

Scarlet Tanager Negative

Summer Tanager Positive

Eastern Towhee Positive

Indigo Bunting Positive Positive

Baltimore Oriole Positive
80 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

conditions permitted few trees to develop and then were kept

open by extensive use of fire by Native Americans. Bird

species that prefer open habitat structure have either largely

disappeared (Greater Prairie Chicken, Bachman’s Sparrow,

Red-cockaded Woodpecker) or shifted to agricultural

habitats. Many of the species that were common in

grasslands prior to European settlement and subsequently

shifted to agricultural habitats are undergoing widespread

population declines as farmland has been converted to

forests or human environments. Great reduction in the

number of grasslands, balds, and barrens has increased the

importance of managing the remaining grasslands with care

to avoid endangering small isolated populations of

invertebrates and some bird species. Prescribed fire is an

effective tool and sometimes the only practical tool for

achieving management goals. Managers can take practical

steps to minimize loss of individuals through the use of

prescribed fire. Any change of fire frequency—either

increasing or decreasing—over a long time period will favor

some plant species over others, which will strongly affect the

habitat structure and associated bird species. In the Central

Hardwoods region widespread declines of bird species that

are associated with open habitats indicates that fire should

be used widely to maintain openings.

Acknowledgments

I thank Jeff Brawn, Chris Bades, Paula Seamon, W. Wilson

Baker, Don Wood, and Dan Wenny for sharing information

and ideas about birds and fire in the Central Hardwoods.

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Continued

Appendix

Bird species of the Central hardwoods landscape. Status: B=breeding, W=winter, and Y=year-round. Species that have

shown statistically significant changes from 1965-1995 from Breeding Bird Survey results. (See text for list of BBS

physiographic regions.)

N physiographic regions

showing significant trends

Common Name Scientific Name Status Positive Negative

Pied-billed Grebe Podilymbus podiceps Y

Eared Grebe Podiceps nigricollis W

American White Pelican Pelecanus erythrorhynchos B

Double-crested Cormorant Phalacrocorax auritus B

Anhinga Anhinga anhinga B

American Bittern Botaurus lentiginosus B

Least Bittern Ixobrychus casmerodius B

Great Blue Heron Ardea herodias B 70

Great Egret Casmerodius albus B

Snowy Egret Egretta thula B

Little Blue Heron Egretta caerulea B

Tricolored Heron Egretta tricolor B

Cattle Egret Bubulcus ibis B

Green Heron Butorides striatus B 02

Black-crowned Night-Heron Nycticorax nycticorax B

Yellow-crowned Night-Heron Nycticorax violaceus B

Canada Goose Branta canadensis Y

Wood Duck Aix sponsa B 40

Green-Winged Teal Anas crecca W

American Black Duck Anas rubripes Y

Mallard Anas platyrhynchos Y 20

Northern Pintail Anas acuta Y

Blue-winged Teal Anas discors B

Northern Shoveler Anas clypeata B

Gadwall Anas strepera W

American Wigeon Anas americana B

Canvasback Aythya valisineria W

Redhead Aythya americana W

Ring-necked Duck Aythya collaris W

Lesser Scaup Aythya affinis W

Common Goldeneye Bucephala clangula W

Bufflehead Bucephala albeola W

Hooded merganser Lophodytes cucullatus B

Common Merganser Mergus merganser W

Ruddy Duck Oxyura jamaicensis W

Black Vulture Coragyps atratus B 10

Turkey Vulture Cathartes aura Y 10

Osprey Pandion haliaetus B

Mississippi Kite Ictinia mississippiensis B

Bald Eagle Haliaeetus leucocephalus W

Northern Harrier Circus cyaneus Y

Sharp-shinned Hawk Accipiter striatus Y 10

Cooper’s Hawk Accipiter cooperii Y 10

Northern Goshawk Accipiter gentilis W

Red-shouldered Hawk Buteo lineatus Y 30

Broad-winged Hawk Buteo platypterus B 01

Red-tailed Hawk Buteo jamaicensis Y 70

Rough-legged Hawk Buteo lagopus W

Golden Eagle Aquila chrysaetos W
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 83

Appendix continued

N physiographic regions

showing significant trends

Common Name Scientific Name Status Positive Negative

Continued

American Kestrel Falco sparverius Y 11

Merlin Falco columbarius W

Gray Partridge Perdix perdix Y

Ring-necked Pheasant Phasianus colchicus Y

Ruffed Grouse Bonasa umbellus Y

Sharp-tailed Grouse Tympanuchus phasianellus Y

Greater Prairie Chicken Tympanuchus cupido Y

Wild Turkey Meleagris gallopavo Y

Northern Bobwhite Colinus virginianus Y

King Rail Rallus elegans B

Virginia Rail Rallus limicola B

Sora Porzana carolina B

Purple Gallinule Porphyrula martinica B

Common Moorhen Gallinula chloropus B

American Coot Fulica americana Y

Piping Plover Charadrius melodus B

Killdeer Charadrius vociferus Y 41

Spotted Sandpiper Actitis macularia B

Upland Sandpiper Bartramia longicauda B

Common Snipe Gallinago gallinago W

American Woodcock Scolopax minor Y

Wilson’s Phalarope Phalaropus tricolor B

Ring-billed Gull Larus delawarensis W

Herring Gull Larus argentatus W

Common Tern Sterna hirundo B

Forster’s Tern Sterna forsteri B

Least Tern Sterna antillarum B

lack Tern Chlidonias niger B

Black Skimmer Rynchops niger B

Rock Dove Columba livia Y

Mourning Dove Zenaida macroura Y 40

Black-billed Cuckoo Coccyzus erythropthalmus B 01

Yellow-billed Cuckoo Coccyzus americanus B 06

Common Barn-Owl Tyto alba Y

Eastern Screech-Owl Otus asio Y

Great Horned Owl Bubo virginianus Y

Barred Owl Strix varia Y

Long-eared Owl Asio otus Y

Short-eared Owl Asio flammeus Y

Northern Saw-whet Owl Aegolius acadicus Y

Common Nighthawk Chordeiles minor Y 02

Chuck-will’s-widow Caprimulgus carolinensis B

Whip-poor-will Caprimulgus vociferus B 02

Chimney Swift Chaetura pelagica B 13

Ruby-throated Hummingbird Archilochus colubris B 01

Belted Kingfisher Ceryle alcyon Y 13

Red-headed Woodpecker Melanerpes erythrocephalus Y

Red-bellied Woodpecker Melanerpes carolinus Y

Yellow-bellied Sapsucker Sphyrapicus varius W

Downy Woodpecker Picoides pubescens Y

Hairy Woodpecker Picoides villosus Y

Red-cockaded Woodpecker Picoides borealis Y
84 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

N physiographic regions

showing significant trends

Common Name Scientific Name Status Positive Negative

Appendix continued

Continued

Northern Flicker Colaptes auratus Y

Pileated Woodpecker Dryocopus pileatus Y

Olive-sided Flycatcher Contopus borealis B

Eastern Wood-Pewee Contopus virens B 05

Acadian Flycatcher Empidonax virescens B 02

Alder Flycatcher Empidonax alnorum B

Willow Flycatcher Empidonax traillii B

Least Flycatcher Empidonax minimus B 01

Eastern Phoebe Sayornis phoebe B 30

Great Crested Flycatcher Myiarchus crinitus B 05

Eastern Kingbird Tyrannus tyrannus B 02

Scissor-tailed Flycatcher Tyrannus forficatus B

Horned Lark Eremophila alpestris Y 04

Purple Martin Progne subis B 12

Tree Swallow Tachycineta bicolor B 10

Northern Rough-winged SwallowStelgidopteryx serripennis B 21

Bank Swallow Riparia riparia B

Cliff Swallow Hirundo pyrrhonota B 10

Barn Swallow Hirundo rustica B 12

Blue Jay Cyanocitta cristata Y

American Crow Corvus brachyrhynchos Y

Fish Crow Corvus ossifragus B

Common Raven Corvus corax Y

Black-capped Chickadee Parus atricapillus Y

Carolina Chickadee Parus carolinensis Y

Tufted Titmouse Parus bicolor Y

Red-breasted Nuthatch Sitta canadensis W

White-breasted Nuthatch Sitta carolinensis Y

Brown-headed Nuthatch Sitta pusilla Y

Brown Creeper Certhie americana W

Carolina Wren Thryothorus ludovicianus Y

Bewick’s Wren Thryomanes bewickii B 01

House Wren Troglodytes aedon B 42

Winter Wren Troglodytes troglodytes W

Marsh Wren Cistothorus palustris B

Golden-crowned Kinglet Regulus satrapa W

Ruby-crowned Kinglet Regulus calendula W

Blue-gray Gnatcatcher Polioptila caerulea B 10

Eastern Bluebird Sialia sialis Y 41

Veery Catharus fuscescens B

Swainson’s Thrush Catharus ustulatus B

Hermit Thrush Catharus guttatus W

Wood Thrush Hylocichla mustelina B 14

American Robin Turdus migratorius Y 70

Gray Catbird Dumetella carolinensis B 24

Northern Mockingbird Mimus polyglottos Y

Brown Thrasher Toxostoma rufum B

Cedar Waxwing Bombycilla cedrorum Y 60

Northern Shrike Lanius excubitor W

Loggerhead Shrike Lanius ludovicianus Y 03

European Starling Sturnus vulgaris Y

White-eyed Vireo Vireo griseus B 13
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 85

N physiographic regions

showing significant trends

Common Name Scientific Name Status Positive Negative

Appendix continued

Continued

Solitary Vireo Vireo solitarius B

Bell’s Vireo Vireo bellii B 01

Yellow-throated Vireo Vireo flavifrons B 01

Warbling Vireo Vireo gilvus B 11

Red-eyed Vireo Vireo olivaceus B 11

Blue-winged Warbler Vermivora pinus B 02

Golden-winged Warbler Vermivora chrysoptera B 01

Nashville Warbler Vermivora ruficapilla B

Northern Parula Parula americana B 10

Yellow Warbler Dendroica petechia B 20

Chestnut-sided Warbler Dendroica pensylvanica B

Magnolia Warbler Dendroica magnolia B

Black-throated Blue Warbler Dendroica caerulescens B

Yellow-rumped Warbler Dendroica coronata W 10

Black-throated Green Warbler Dendroica virens B 10

Blackburnian Warbler Dendroica fusca B

Yellow-throated Warbler Dendroica dominica B 21

Pine Warbler Dendroica pinus B 20

Prairie Warbler Dendroica discolor B 06

Cerulean Warbler Dendroica cerulea B 03

Black-and-white Warbler Mniotilta varia B 04

American Redstart Setophaga ruticilla B 04

Prothonotary Warbler Protonotaria citrea B 01

Worm-eating Warbler Helmitheros vermivorus B

Swainson’s Warbler Limnothlypis swainsonii B

Ovenbird Seiurus aurocapillus B

Louisiana Waterthrush Seiurus motacilla B

Common Yellowthroat Geothlypis trichas B 02

Hooded Warbler Wilsonia citrina B

Kentucky Warbler Oporornis formosus B 01

Canada Warbler Wilsonia canadensis B

Yellow-breasted Chat Icteria virens B 07

Summer Tanager Piranga rubra B 02

Scarlet Tanager Piranga olivacea B 31

Northern Cardinal Cardinalis cardinalis Y

Rose-breasted Grosbeak Pheucticus ludovicianus B 21

Blue Grosbeak Guiraca caerulea B

Indigo Bunting Passerina cyanea B 07

Painted Bunting Passerina ciris B

Dickcissel Spiza americana B 02

Rufous-sided Towhee Pipilo erythrophthalmus Y 09

Bachman’s Sparrow Aimophila aestivalis B

American Tree Sparrow Spizella arborea W

Chipping Sparrow Spizella passerina B 22

Field Sparrow Spizella pusilla Y 0 10

Vesper Sparrow Pooecetes gramineus B 03

Lark Sparrow Chondestes grammacus B 01

Savannah Sparrow Passerculus sandwichensis B

Grasshopper Sparrow Ammodramus savannarum B 06

Henslow’s Sparrow Ammodramus henslowii B

Le Conte’s Sparrow Ammodramus leconteii W

Fox Sparrow Passerella iliaca W
86 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

N physiographic regions

showing significant trends

Common Name Scientific Name Status Positive Negative

Appendix continued

Song Sparrow Melospiza melodia Y

Lincoln’s Sparrow Melospiza lincolnii W 10

Swamp Sparrow Melospiza georgiana Y

White-throated Sparrow Zonotrichia albicollis W

White-crowned Sparrow Zonotrichia leucophrys W

Dark-eyed Junco Junco hyemalis W

Lapland Longspur Calcarius lapponicus W

Snow Bunting Plectrophenax nivalis W

Bobolink Dolichonyx oryzivorus B 11

Red-winged Blackbird Agelaius phoeniceus Y 06

Eastern Meadowlark Sturnella magna Y 09

Western Meadowlark Sturnella neglecta B

Yellow-headed Blackbird Xanthocephalus xanthocephalus B

Rusty Blackbird Euphagus carolinus W

Brewer’s Blackbird Euphagus cyanocephalus B

Common Grackle Quiscalus quiscula Y

Brown-headed Cowbird Molothrus ater Y 03

Orchard Oriole Icterus spurius B 13

Northern Oriole Icterus galbula B 13

Purple Finch Carpodacus purpureus W

House Finch Carpodacus mexicanus Y

Pine Siskin Carduelis pinus W

American Goldfinch Carduelis tristis Y 01

Evening Grosbeak Coccothraustes vespertinus W

House Sparrow Passer domesticus Y

Eurasian Tree Sparrow Passer montanus Y
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 87

Despite the widespread use of fire as a management tool in

forested landscapes, we know surprisingly little about its

effects on streams and their biota. Forest management of

this type may affect light, temperature, nutrient, sediment,

and hydrologic regimes within stream channels, which in turn

may affect the abundance and diversity of aquatic microbes,

plants, and animals. In cooperation with the USDA Forest

Service / National Forests in Alabama, in 1994 my graduate

students and I began a long-term experimental study of the

effects of forest thinning and prescribed burning on stream

water quality within the Talladega National Forest (Shoal

Creek District), Alabama. Streams in six small watersheds

have been monitored seasonally; three of the streams are

being exposed to understory thinning and infrequent, lowintensity burning (i.e., once every 2 years), and the three

remaining streams drain watersheds that have been left

intact as reference sites. Study watersheds lie in close

proximity to each other and have similar drainage areas,

geomorphology, parent geology, and soils. Streams are

small (1st-order channels, < 2 m wide), heavily shaded (>

85% riparian cover), and are well oxygenated (> 7 ppm).

Particular emphasis has been placed on the response of

benthic invertebrates (mostly aquatic insects) to

management, although we also have studied physicalchemical water quality attributes and stream periphyton

(attached algae).

Our analyses to date revealed no major differences in water

chemistry between managed and reference streams, with

both groups of streams showing uniform pH (slightly acidic),

and low alkalinity, conductivity, total suspended solids, and

dissolved nutrients. Periphyton abundance was higher in

spring and summer (vs. winter), and also was generally

higher in managed streams. All streams showed high benthic

macroinvertebrate species richness (> 125 species per

stream). Preliminary analysis from spring and summer 1995

samples suggests that habitat-specific richness was higher

in reference than in managed streams, especially in spring.

However, the two groups of streams showed similar total

invertebrate total density, species diversity, EPT richness

(i.e., number of mayfly, stonefly, & caddisfly species), and

species dominance. Although these data are preliminary,

they are compelling in that they suggest thinning and burning

and associated environmental changes (e.g., increases in

light levels reaching streams, short-term but biologically

significant nutrient releases, etc.) may have positive effects

on the abundance and species composition of stream

periphyton, but only minor effects on stream invertebrate

diversity.

Stream Mayflies, Stoneflies, and Caddisflies:

Do They Care About Controlled Burning in the Forest?

Jack W. Feminella1

1

Associate Professor, Department of Biological Sciences,

Auburn University, 331 Funchess Hall, Auburn, AL 36849-5407,

email: feminjw@mail.auburn.edu
88 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

In 1990, I initiated insect surveys for the Kentucky Field

Office of The Nature Conservancy (KYFO) on several of the

highest quality oak/pine barrens and sandstone/limestone

glades left in the state. These plant communities are known

to contain many state and globally rare plant species found

only on these remnant natural areas. Therefore, it was

thought that they might also contain remnant-requiring

insects. Given that these plant communities are dominated

by grasses and fire-dependent plant species, fire was an

important part of their history and is used widely in their

Fire Bugs

Jim Bess1

13501 South 750, West Wanatah, IN 46390, ph.: 219-733-2947,

email: jabess@mail.netnitco.net

current management and restoration. The KYFO was

primarily interested in identifying those species restricted in

occurrence to barrens and glade remnants which might also

be sensitive to the short-term effects of prescribed burning.

Our studies have focused on the three major herbivorous,

grassland insect orders: the Homoptera (cicadas,

leafhoppers, aphids and their relatives), the Lepidoptera

(butterflies and moths) and Orthoptera (grasshoppers,

katydids, crickets, mantids and stick insects). The results of

my survey work have identified a large number of potentially

remnant-requiring insects (roughly 25 percent of the total),

several of which are new to the state. On going survey work

is continuing to identify new species and we are focusing our

attention on an even broader assortment of insect taxa.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 89

Abstract

The black bear (Ursus americanus) is a species of interest

for wildlife and forest managers, landowners, hunters, and

the public. Because of their overall adaptability to changing

habitat conditions, wide-ranging nature, and omnivorous

habits, black bears are readily accommodated in a forest

management program. A well-managed forest that provides

adequate opportunities for movements, foraging, denning,

and cover has better potential to satisfy the needs of bears

than an unmanaged forest. Food supplies should be

abundant, stable, and diverse. Habitats that harbor natural

dens should be conserved, known den sites should be

protected, and creation of potential den habitat and den sites

should be considered in forest habitat treatments. In

addition, natural escape cover should be maintained, and

opportunities should be undertaken to create or enhance

cover for bedding, escape, and dispersal. Although bear

habitat management has traditionally been associated with

timber management, the use of prescribed fire holds promise

as an effective tool for enhancement of black bear habitat in

the central hardwood landscape. In this paper, various

aspects of black bear habitat ecology are reviewed, and the

use of prescribed fire alone or in conjunction with other

silvicultural techniques to maintain, enhance, and restore

habitat conditions for black bears is explored. Evaluation of

prescribed fire behavior and activity, habitat responses to fire

under various conditions, and associated implications for

bears could be useful in developing “best fire management

practices” in bear habitat. Prescribed fire should be used in

an adaptive management approach. This entails not only the

application of fire, but also the diligent monitoring of habitat

and wildlife responses, and fine-tuning future actions. Topics

of investigation concerning bears and fire are listed. Public

education programs conveying the ecological effects and

benefits of prescribed fire for habitat, wildlife, and humans,

and the development of public/private partnerships for

habitat management are recommended.

Background

The black bear is a species of management interest in most

areas of its range. Humans hold widely divergent viewpoints

regarding bears. Black bears are characterized as big game

animals, links to cultural heritage, food sources, dangerous

animals, tourist attractions, photographic subjects, spiritual

beings, indicators of healthy ecosystems, symbols of

wilderness, pawns for environmental activism, agricultural

pests, and as a threat to resource extraction and economic

development programs. Because of the importance of bears

in the lives of many humans, and the need to satisfy

legislative, regulatory, or other obligations to properly

manage wildlife and their habitats, wildlife and land

managers seek to enact management practices beneficial to

bears and humans. Responsible bear management includes

the recognition that carrying capacity has biological and

socioeconomic aspects. Both must be considered for their

relevance to providing adequate resources for bears,

reducing human/bear conflicts, and fostering public

acceptance of bears. Bear damage complaints in agriculture

and garbage usually increase when shortages of natural

foods occur, which provides additional justification for

establishing and maintaining suitable forest conditions for

bears. This is particularly necessary in fragmented habitat

where bears are surrounded by more agriculture and

development than forest. Bear habitat management should

focus on providing suitable habitat and habitat linkages,

abundant natural food supplies, denning sites, escape cover,

and human access management. The purpose of this paper

is to highlight some aspects of black bear habitat ecology

and to explore the use of prescribed fire to enhance bear

habitat in the central hardwood landscape. The concept of

prescribed fire as used in this paper is the application of

human ignited and controlled fire to a site, according to

approved techniques, within specified limits of the physical

conditions of air, soil, and vegetation, in order to effect a

desired change in site conditions, for the purpose of

achieving previously defined goals and objectives.

Habitat

Black bears are adaptable, opportunistic, and highly mobile

animals. Bears can move great distances in response to

changing weather, food supplies, habitat conditions,

disturbance, and internal population pressures. Movements

and home range size vary greatly depending on age, sex,

reproductive status, population density, and habitat quality.

Males typically range farther than females. Sizes of bear

home ranges in the Cherokee National Forest in east

Tennessee, as determined through analysis of radiotelemetry

data, varied from 20 to 192 square kilometers for males and

4 to 23 square kilometers for females (Villarubia 1982, Garris

1983, Clevenger 1986). Such mobility simplifies habitat

management for bears in some respects, because

management actions can be directed beyond the microsite to

the forest or landscape scale. Conservation or creation of

habitat linkages between disjunct tracts of forest can help

provide corridors for bear movements in search of food,

dens, and mates; juvenile dispersal; and also facilitate bear

population expansion.

Although black bears are creatures of the forest, they do not

require wilderness or old growth habitats. However, both of

these habitat characteristics can be useful components of

the overall bear landscape. Black bears can survive and

coexist with human inhabitation nearby if the bears are

afforded a forest that satisfies their life requisites. The use of

Black Bear Ecology and the Use of Prescribed Fire to Enhance Bear Habitat

Keith M. Weaver 1

1

Refuge Manager, U.S. Fish and Wildlife Service, Nulhegan

Basin Division, Silvio O. Conte NFWR, P.O. Box 427, Island

Pond, VT 05846.
90 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

prescribed fire can aid in establishment of desired

regeneration. It also can be used for site preparation in

reforestation programs that will expand forested habitat and

provide an additional buffer from human encroachment into

bear habitat. The greater the quantity, but perhaps more

important, the quality of bear habitat, the less chance for

conflicts between humans and bears.

The key to improving the quality of bear habitat is to provide

habitat diversity. Within the forest, a mosaic of habitats or

habitat conditions (e.g., species composition, age class,

structure, successional stage, openings, understory) holds

the best potential for the satisfaction of bear life

requirements. The value of managing a mixed pine-oakhickory component within the central hardwood landscape

should not be overlooked for its benefit for bears. A diverse

forest may be better adapted to withstand the ravages of

disease, insect invasions, or catastrophic events. Therefore,

a properly managed forest likely will provide better bear

habitat potential than an unmanaged forest. Indeed, the

outcome of modeling timber harvest effects on black bear

population dynamics in the southern Appalachian Mountains

indicated that forests with carefully designed timber

management regimes would yield greater food abundance

and stability than unmanaged forests (Brody and Stone

1987). It is important to recognize that a decision not to

perform habitat manipulation has its own consequences that

also must be evaluated for their impact on bears.

The credibility and compatibility of forest management in

providing for bear habitat needs was demonstrated by the

U.S. Fish and Wildlife Service through promulgation of a

special rule in the Threatened listing for the Louisiana black

bear (U. a. luteolus), which states, in part, that the provisions

of the Federal listing “... shall not prohibit effects incidental to

normal forest management activities within the historic range

of the Louisiana black bear except for activities causing

damage to or loss of den trees, den tree sites or candidate

trees” (Federal Register 7 January 1992, 57(4) sub-sect.

17.40, i(2), pp. 594-595). Normal forest management

activities were defined in the rule as “... activities that support

a sustained yield of timber products and wildlife habitats,

thereby maintaining forestland conditions in occupied

habitat.” Although the range of the Louisiana bear is distant

from the central hardwood landscape, the premise is

identical; responsible forest management is good bear

habitat management.

Traditionally, forest management for black bears has been a

function of timber management. Treatments such as

clearcut, patch clearcut, shelterwood and irregular

shelterwood, and thinning are useful methods for creating,

improving, and maintaining desirable bear habitat. However,

the ability to manage forests with conventional silvicultural

techniques has been hindered because of increasing public

resentment to timber harvest, increased environmental

regulation, and decreasing budgets. Therefore, the use of

prescribed fire may provide a complementary habitat

management technique for bears. Prescribed fire also may

be a useful tool to enhance bear habitat in areas where

timber harvest is impractical or prohibited. Ideally, the

integration of prescribed fire with other silvicultural

treatments affords the greatest potential for maintenance

and establishment of suitable bear habitat conditions. A

comprehensive prescribed fire program also should include

hazard fuel reduction burns that serve to reduce the

occurrence or severity of wildfires that could destroy bear

habitat and negate beneficial habitat management programs.

The practice of managing forests, landscapes, or wildlife

populations implies making choices that entail potential risks

and benefits, and advantages and disadvantages for target

and nontarget sites, species, and humans. Therefore, it is

imperative that clear management goals and objectives are

defined and approved, appropriate methods employed,

outcomes evaluated for success or failure, and modifications

for future achievement enacted. Prescribed fire use must not

be relegated to the “meet the target” syndrome. Additionally,

a forest management program should not be expected to

provide for every wildlife or human use, on every acre, in

every management compartment, or in every decade. Some

sites are better suited to a certain management purpose than

others, and there are practical limitations to the degree of

manipulation that can be achieved or the extent of attention

that can be devoted on a site during any given period.

Tradeoffs will occur and management choices must be made

that will accommodate the greatest suite of needs and

interests within applicable laws, policy, funding, technology,

and practicality. These choices should be based on the best

available technical information; however, when developing

new practices, guiding methodologies are often limited, and

sometimes the intuition and experience of managers must be

relied on to determine the prudent course of action.

Foods

Nutrition is the foundation for the survival and welfare of

individual bears and the dynamics of bear populations (e.g.,

Rogers 1976, 1987, 1993). Therefore, it is critical that forest

management practices for black bears ensure a variety of

stable and abundant food supplies throughout the year. This

can be accomplished by providing habitat diversity; a mosaic

of habitat types, stand ages, and conditions; high yields of

soft and hard mast; herbaceous foods; and decomposing

wood that harbors insect and other invertebrate life. Forested

habitat, forest openings, edge habitats, riparian borders, and

crop fields within and adjacent to forested habitat provide

feeding opportunities for bears. Black bears are excellent

tree climbers and are able to forage in all vertical strata of

the forest; therefore, the impacts of habitat management

actions on food production in the overstory, midstory, and

understory should be considered. Black bears are wideranging, omnivorous, and opportunistic in their feeding

habits. Thus they are able to capitalize on a wide range of

conditions in their search for food. Food “preferences”

(Bacon and Burghardt 1983) are difficult to determine, but

bear foraging behavior and food consumption are influenced

by: season, food abundance and accessibility, proximity of

other foraging bears, bear densities, human activity in bear

habitat, physiological condition (e.g., pre and post denning),

and reproductive status.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 91

Hard Mast

The use of the term hard mast in this paper refers

collectively to the fruit of oaks (Quercus spp.), beechnuts

(Fagus grandifolia), and hickories and pecans (Carya spp.).

In the fall and early winter prior to denning, bears in the

eastern deciduous forest feed heavily on hard mast,

particularly acorns. They largely depend on these foods to

build sufficient fat reserves to survive the period of winter

dormancy (Beeman and Pelton 1980, Eagle and Pelton

1983, Clark and others 1987, Rogers and Allen 1987). Bears

forage for hard mast in tree crowns prior to nut drop, and on

the ground post-drop. The abundance and availability of hard

mast affect black bears at the individual and population

levels. The availability of hard mast, particularly where there

are few other foods, has been documented to affect

population growth (mortality, natality, interbirth interval), bear

numbers, fall feeding movements, and denning chronology

(Pelton 1985, 1989; McLaughlin and others 1994;

McLaughlin 1998; Schooley and others 1994). Such effects

of hard mast on black bear survival and population dynamics

in the Southern Appalachians have been modeled (Brody

and Stone 1987, McLean and Pelton 1994).

Hard mast production should be an important consideration

in bear habitat management in the central hardwood

landscape. Where feasible, hard mast producing species,

particularly oaks, should be encouraged as a major bear

forest habitat component in silvicultural and prescribed fire

treatments. Long timber rotations (75 to 100 years) will favor

hard mast production, and are compatible with peak acorn

yield potential (Pelton 1985, Hillman and Yow 1986, Brody

and Stone 1987). Treatments should be directed toward

maintaining diversity of hard mast species, high mast yields,

and perpetuating desired species in future stands. A notable

problem in upland hardwood management has been the

difficulty in regenerating oak stands on high quality sites

following harvest. Thus, many oak-dominated stands have

been converted to stands of less desirable species in terms

of hard mast such as yellow poplar (Liriodendron tulipifera),

red maple (Acer rubrum), and sweetgum (Liquidambar

styraciflua) (O’Hara 1986, Kays and others 1988). If this

conversion occurs on a wide scale it could adversely affect

bear populations. However, upland oaks exhibit

morhpological and physiological characters that suggest

greater fire adaptation than some of their competing species

including: thick bark, vigorous sprouting ability, deep roots,

rot resistance following scarring, xeromorphic leaves, and

greater hardiness to drought and nutrient-poor sites (Abrams

1992). Historically, oak domination on central hardwood sites

has been in response to a fire regime (Abrams 1992).

Advance regeneration in the understory is a prerequisite for

reestablishment of an oak-dominated stand subsequent to

harvest. Van Lear and Waldrop (1989) suggested that

repeated understory burns in summer and winter during an

extended period (e.g., 5 to 20 years) prior to harvest could

be successful in establishing adequate oak advance

regeneration. Recently, compelling evidence has been

presented that the proper timing, intensity, and periodicity of

prescribed fire applied 3 to 5 years following shelterwood

cutting in stands with advance oak regeneration will favor

oak reproduction and establishment on productive upland

sites. The concept, development, and applications of this

technique as well as instructions for its implementation are

presented by Brose and Van Lear (1998, 1999) and Brose

and others (1999a,b). The benefits and drawbacks of

prescribed burning in oak-dominated shelterwood stands in

different seasons also are discussed by Brose and Van Lear

(1998, 1999). The effects of prescribed burning on hard mast

production were discussed by Harlow and Van Lear (1989).

They commented that hot surface fires in the fall could

readily consume acorns in the duff layer, but that low

intensity surface fires could serve to expose recently fallen

acorns without substantial loss of mast crops. Consumption

of heavy leaf litter by fire could aid in the successful location

and caching of acorns by blue jays (Cyanocitta cristata) and

squirrels (Sciurus spp.), which could promote oak

regeneration (Galford and others 1988). They also indicated

winter burning might reduce acorn predation by weevils, thus

increasing their availability for wildlife.

Squawroot (Conopholis americana), a saprophyte

associated with oaks, is an important component of spring

diets of bears following den emergence. Squawroot provides

nutrition for bears at a period when soft mast production is

not yet available. The perpetuation of an oak component in

the forest would hopefully enable continued availability of

squawroot. The ecology of oaks, squawroot, and fire

warrants investigation.

Soft mast

Although hard mast crops are important for black bears, they

only are seasonally present, and abundant yields are

sporadic. Therefore, other foods also must be available

throughout the year. In addition, during hard mast failures,

alternate food sources are of great importance. Soft mast

produced by woody and herbaceous species is a vital food

source that also should be managed as a viable component

of bear habitat. Use of the term soft mast in this paper refers

to fruits with fleshy exteriors such as berries, drupes, and

pomes (Harlow and Van Lear 1989). In the central hardwood

landscape, bears consume the fruits of a wide variety of

plants such as: blackberries (Rubus spp.), blueberries

(Vaccinium spp.), huckleberries (Gaylussacia spp.),

serviceberry (Amelanchier spp.), pokeberry (Phytolacca

americana), elderberry (Sambucus canadensis), devil’s

walking stick (Aralia spinosa), flowering dogwood (Cornus

florida), paw-paw (Asimina triloba), sassafras (Sassafras

albidum), Ilex spp., hawthorn (Crataegus spp.), black cherry

(Prunus serotina), black gum (Nyssa sylvatica), and

persimmon (Diospyros virginiana). Growth of most of these

plants are stimulated by increased light penetration to the

forest floor from canopy openings that result from timber

harvest, construction of wildlife openings, creation of interior

forest trails, and wind or ice damage. Silvicultural and

prescribed fire treatments that promote establishment,

diversification, proliferation, and perpetuation of soft mast as

a forest component in conjunction with hard mast are highly

encouraged. Soft mast production peaked 3 to 5 years after

clearcuts, shelterwood cuts, and group selection treatments

in mixed pine-hardwood stands in the Oauchita Mountains of
92 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Arkansas and Oklahoma (Perry and others 1988). However,

it was noted that without additional treatment, such as

burning or thinning, soft mast production was expected to

decline significantly as canopies in the harvested areas

closed. Fruit producing vines such as grape (Vitis spp.) and

greenbrier (Smilax spp.) are also readily eaten by bears;

their growth should be encouraged. Heavy canopies of vines

(e.g., grape slicks) generally are not compatible with oak

regeneration, but if these growths are managed for fruit

production (versus mainly vegetative growth), they add a

valuable component of diversity for bear foods, and can be

important to bears during shortages of hard mast.

The effects of prescribed fire on soft mast production in

hardwood stands has received little attention in the literature.

A literature review of the effects of prescribed fire on soft

mast production in various pine forest types in the Southeast

was presented by Harlow and Van Lear (1989). Many soft

mast producing species considered in this paper decreased

fruit production in the year following burning; however,

fruiting in flowering dogwood was increased in the next year

after burning. Studies reviewed indicated that fruiting

increased 2 to 5 years following burning in species that occur

in the central hardwoods such as blueberries and

huckleberries, blackberries, persimmon, chokeberry (Sorbus

arbutifolius), muscadine grape (V. rotundifolia), Viburnum

spp., yaupon (I. vomitoria) and plum (Prunus umbellata).

Harlow and Van Lear (1989) suggested staggered burning

cycles of 3 to 5 years. Juxtaposition of burned and unburned

areas to promote stable and abundant soft mast crops also

has been recommended (Landers 1987, Harlow and Van

Lear 1989). Burning rotations of 3 to 10 years were

recommended for soft mast production for bears in pinehardwood stands in coastal North Carolina (Hamilton 1981)

and pine dominated habitat in the panhandle of Florida by

(Stratman 1998).

Herbaceous Foods

Bears forage in the forest understory, forest openings, and

edge habitats for herbaceous plants. The succulent sprouts

of various plants stimulated by prescribed fire can provide

feeding opportunities in the understory. Pack and others

(1988) documented a 31 percent increase in herbaceous

forbs, grasses, and sedges following late winter-early spring

burns in oak-hickory stands in West Virginia that had

previously been thinned. Treatments involving thinning alone

or thinning following burning did not produce similar results.

To prevent risk of fire injury to the residual stand, they

recommended burning only after logging slash had

decomposed to the point where it would not constitute a

major fuel component. Masters and others (1993), studying

timber harvest and fire effects in pine-oak-hickory stands in

the Ouachita Mountains of Oklahoma, reported that fire

could reduce accumulations of litter and standing dead

herbaceous vegetation that impeded new herbaceous

growth. The combination of timber harvest and burning

increased light penetration to the forest floor, resulted in

increased soil temperatures and nitrogen availability, which,

in turn, stimulated earlier and greater herbaceous growth

and production.

Establishment of a system of widely dispersed forest

openings, perhaps <1 ha in size, maintained in various age

and species compositions, would provide additional

diversification of bear foraging opportunities. Abandoned log

loading sites and old fields can be converted to wildlife

openings and managed to produce herbaceous vegetation.

Old trails or roads not used for transportation routes within

the forest can be converted and maintained as linear wildlife

openings (Hillman and Yow 1986). Openings could even be

created in marginal forest stands using a hot fire that would

kill woody vegetation. Annual burning likely will be necessary

to stimulate high seed production and maintain herbaceous

conditions. Disking or mowing on a 1 to 3 year rotation can

be used to maintain openings in early successional stages.

However, prescribed fire could provide an alternate method

for rejuvenation of herbaceous growth and to setback woody

succession. Use of prescribed fire can eliminate the need of

accessing remote openings with machinery, and reduce

costs of maintaining and operating tractors and implements.

Openings can be enhanced with periodic sowing of food

plants such as clovers (Trifolium spp.), wheat, rye, chufa

(Cyperus esculentus), brown-top millet (Panicum ramosum),

or bahia grass (Paspalum notatum). Prescribed fire can aid

site preparation for initial or subsequent seedings. Edge

plants such as blackberries, pokeweed, elderberry, and

devil’s walking stick that proliferate in and around forest

openings further increase habitat diversity and feeding

opportunities. These openings also provide habitat for a host

of other wildlife species. Prescribed fire management is also

indicated for vegetation management in rights-of-way, and

for the perpetuation and management of grassy balds in the

Southern Appalachians. Resultant vegetation and soft mast

within and along the edges of these treatment areas serve to

increase bear feeding opportunities.

Invertebrates

Invertebrates such as beetles, ants, termites, and their eggs

and larva are protein-rich foods for bears. Such food items

often are found inside or underneath decaying trees and

logs. Decayed logs that are rolled over or torn apart in

search of insects are a common sight in most areas of bear

activity. These logs typically are remnants of logging slash or

fallen dead trees. Black bears consume insects during most

of their non-denning period. Bears also feed on wild honey

bees and their honey that are found in standing hollow dead

trees, and in cavities in living trees. Retention of standing

and fallen snags, and logging slash is encouraged to

provide invertebrate food sources. When planning a

prescribed fire program with bear habitat enhancement in

mind (versus strictly hazard fuel reduction, for example),

such food shelters for bears should be favored. Some areas

typically targeted for slash reduction burns should be left

unburned, or at a minimum, some slash should be protected

from fire near the forest edge. Slash left solely for bear

foraging opportunities should not be piled. When using

prescribed fire in the understory, some fallen logs and

standing snags should be protected from fire when and

where feasible.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 93

Denning Sites

In the central hardwood landscape, bears may occupy a

variety of winter den types including elevated tree cavities,

cavities in the base of a hollow tree or under boulders;

excavated cavities, hollow logs, brushpiles, or ground nests.

Typical bear den trees are: living (although dead trees also

are used), large (dbh >90 cm), and old (>150 years).

Entrances to elevated cavities may range from 5 to 30

meters above ground. Entrances to den cavities are either

through a broken top or a side hole caused by a limb break.

Cavities in the base of hollow trees are often initially caused

by bark injury (including fire injury) that allowed entry of fungi

or other agents that eventually resulted in cavity creation.

Potential den tree species in the central hardwoods would

include: northern red oak (Q. Rubra), chestnut oak (Q.

Prinus), white oak (Q. Alba), yellow poplar, red maple, yellow

birch (B. alleghaniensis), white pine (Pinus strobus), and

eastern hemlock (Tsuga canadensis). Excavations usually

occur on side hills under a blowdown, root cavity, hollow tree

base, log, or stump. Brushpile dens typically are found in

logging slash or sites of extensive wind throw. Ground nests

are usually located in dense thickets of woody vegetation

such as rhododendron (Rhododendron maximum), mountain

laurel (Kalmia latifolia), or in regenerating clearcuts.

Recommendations for timber management for black bears in

the southern Appalachian Mountains suggest that potential

den trees should be reserved from logging, and that a

minimum of 5 to10 percent of the forest be maintained in age

classes of 250 years or older to ensure adequate supplies of

den trees (Pelton 1985, Hillman and Yow 1986). Retention of

individual large trees that contain elevated cavities, and large

trees with heart rot or wind, lightning, or fire injuries that

could develop cavities for dens, is also encouraged during

harvest operations. Preferably, such trees should be left in

small reserve clumps to help prevent wind throw, particularly

in clearcuts or shelterwood cuts. Ideally, pre-burn scouting

should be performed to identify potential bear den trees, and

these should be protected from fire by clearing woody debris

and other fuels from around the base (particularly in the case

of standing dead snags). Similarly, downed hollow logs large

enough to accommodate a denning bear (approximately 50

cm in diameter) should be protected from fire consumption.

When silvicultural treatments are to be followed by

prescribed burns, the logging operators should be instructed

to clear tops and other slash away from reserved trees to

protect them from flareup during the burn.

Ground denning (excavations, brushpiles, ground nests)

opportunities are increased following most silvicultural

treatments. Practical methods to provide ground denning

sites include promoting thick regeneration and scattered

piling of felled tops to create potential brushpile den sites.

Like old growth habitat, early successional habitat also

should be managed as a viable bear habitat component. In

forests where few opportunities exist for tree denning (either

due to age class structure, tree species composition, or past

management practices), regeneration resulting from timber

harvest could be crucial habitat for denning bears. Under

such circumstances, a forest managed with consideration for

tree and ground denning sites for bears will provide more

varied denning potential than an unmanaged forest.

However, such logic notwithstanding, the conservation of

dense rhododendron and laurel thickets, which is contrary to

promoting timber regeneration, also will provide ground

denning habitat for bears. An integrated approach of

providing early successional habitat (regeneration) and

maintaining dense shrub thickets would diversify and expand

availability of ground denning habitat.

Although no studies in the central hardwood region have

dealt with the issue of prescribed fires and denning bears, a

study in coastal North Carolina (Lombardo 1993) and two

studies in Florida (Land 1994, Stratman 1998) documented

abandonment of ground nests by bears, including a female

with cubs, during prescribed burns. The bears in the

Lombardo study subsequently redenned; such information

was not reported in the Florida studies. Conversely, on the

Appalachicola National Forest in Florida, denning bears were

not disturbed from ground nests during prescribed burn

operations (Seibert 1993). These reports serve to remind the

manager that prescribed burns must be undertaken with a

clear recognition and understanding of the potential risks and

benefits to the targeted habitat or wildlife component, both

on the site-specific versus forest scale, and on the individual

animal versus population scale. Burn units must be properly

laid out and of manageable size to allow for responsible

protection of special resources and to minimize potential

adverse effects to the physical environment, habitat, wildlife,

or humans.

Escape Cover

Large expanses of contiguous forest or mountainous,

inaccessible forested terrain serve to insulate bears from

human disturbance. However, as forests become smaller

and more fragmented, and as human encroachment and

disturbance in bear habitat increases, escape cover will

become even more vital to insulate bears from human

activities. Escape cover is recognized as an important

component of bear habitat (Pelton 1985, Rogers and Allen

1987, McLaughlin and others 1988). Black bears can thrive

in close proximity to humans if afforded areas of retreat that

ensure little chance of close contact or visual encounters

with humans (Pelton 1982). Areas of dense or impenetrable

vegetation that limits visibility and hinders travel by human

and dogs, such as rhododendron thickets and regeneration

following clearcuts and shelterwood cuts, can provide high

quality shelter for daybeds and escape cover for bears.

Because dense stands of rhododendron and mountain laurel

are not conducive to oak regeneration, this is another

situation of tradeoffs, one that pits the need for regeneration

versus escape cover.

The utility of large trees as bear escape cover should not be

overlooked. Black bears evolved behaviorally and

morphologically as tree climbers, and are well-adapted for

arboreal escape (Herrero 1972). The largest tree in the area

is often chosen as a resting or bedding spot by females with

cubs. The female can tree the cubs when danger is

perceived, and may accompany them up the tree, protect the
94 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

base of the tree, or flee the area to distract intruders. Bears

also will use large trees for resting. Pockets of old growth

forest can ensure the availability of large trees for escape

and resting, but ideally, scattered large trees should be

available throughout the forest to provide the greatest

opportunity for tree escapes. Just as den trees and

candidate den trees can be reserved from timber harvest

and protected from damage during prescribed burns, so can

escape trees. Such trees actually may serve multiple

purposes as denning, escape, and resting sites.

Roads

Considerable attention has been given to the potential

effects of roads on bears in the Southern Appalachians (Carr

and Pelton 1984, Hillman and Yow 1986, Brody and Pelton

1989, Beringer and others 1990). The impacts of roads on

bears are determined primarily by road structure, location,

traffic characteristics, and timing of road use. Roads no

longer in service and without management purpose should

be closed and reforested, or seeded to serve as linear

wildlife openings (Hillman and Yow 1986, Reed and others

1996). However, existing roads can have a useful function in

a comprehensive fire management program. Forest roads

can serve as firebreaks, primary ignition lines, and escape

routes during prescribed burn operations; avenues into

treated habitat for monitoring and research of the effects of

prescribed fire programs on habitat and wildlife; and as

access for wildfire suppression operations for the protection

of bear habitat. Roads can be gated to reduce vehicular

access into bear habitat, maintained as linear wildlife

openings, and prescribed burned to encourage bear food

plants within and adjacent to the roadbed, and still allow

access for management or emergency purposes.

Summary

Black bears are not obligatory inhabitants of old growth

forest or wilderness areas. Forests managed according to

the basic concepts discussed in this paper have the potential

for providing black bear habitat that is superior to

unmanaged forests. Forest and wildlife managers should not

be restricted in the methods and tools available to

accomplish bear habitat management. Rarely is there only

one “right” way to achieve a desired future forest condition.

Management actions must be driven by overall goals and

objectives. Prescriptive criteria should be formulated to effect

burn conditions that will achieve management objectives for

the burn unit, and such criteria must be strictly adhered to in

“go-no go” decision making. Firing techniques must be

appropriately selected and applied to meet burn objectives.

Burns must be adequately monitored from ignition to

completion to record fire behavior and intensity, and to

ensure prescriptive criteria are being satisfied. Burn units

must be of manageable size to minimize chance of fire

escape, and to have reasonable opportunity to suppress the

fire should prescriptive limits be exceeded, or if adverse

conditions or circumstances are encountered. Burn plans

must define methods to measure whether burns achieved

specified burn objectives or not and why. Fire management

plans must include provisions for monitoring of both first

order fire effects and longer term impacts on habitat and

associated wildlife responses. Results of fire management

actions must be evaluated to ensure goals and objectives

indeed are being accomplished, and future actions modified

as necessary in an adaptive management fashion. Through

such efforts, “best fire management practices” can be

developed that will guide future management actions.

Black bear habitat and landscape management in the central

hardwoods, including the use of prescribed fire and wildfire

suppression, should be directed toward: (1) forest

conservation; (2) creating and maintaining habitat diversity;

(3) providing stable, diverse, and widely available food

supplies throughout the year; (4) maintaining and increasing

suitable denning sites and habitat; (5) providing escape

cover; (6) conservation, enhancement, and establishment of

habitat linkages; (7) reforestation programs; and (8) human

access management. Responsible forest management can

accommodate the needs of consumers, recreationists,

residents, visitors, and bears. Bears and humans can coexist

if habitat is available that satisfies the life requirements of

bears, and if bears do not unduly interfere with human

objectives for use of the same landscape.

As with any wildlife population, the objectives and attitudes

of land owners, land managers, resource users, legislators,

and the general public will determine if bears are considered

a positive or negative benefit, and ultimately, if bears can

survive. Expanding public awareness about the ecology and

management of bears, and the utility and benefits of

prescribed fire and other silvicultural techniques should

encourage acceptance of agency and private programs

designed to conserve, manage, and restore bear

populations. Public education should include the concept of

maintaining functional ecosystems, watersheds, and

landscapes as the basis for conserving a unique wildlife

heritage. A considerable amount of the wildlife habitat in the

eastern United States is in private ownership. Therefore, it is

crucial that private landowners also develop a vested interest

in black bear conservation and management to ensure longterm welfare of the species. Technical advice and assistance

with prescribed fire management of wildlife habitat should be

provided to private landowners by federal and state

agencies. Effective partnerships among agencies,

universities, non-governmental organizations, private

enterprise, and the public will aid in information transfer,

cooperative programs, and increased funding, with the end

result being increased fire management success for the

benefit of bears, other wildlife, and humans in the central

hardwood landscape.

Future topics for study concerning the relationship of bears,

fire, and people in the central hardwood landscape include:

(1) determining the most effective integration of prescribed

fire and other silvicultural techniques to produce a productive

and diverse forest; (2) effects of prescribed fire on hard

mast, soft mast, squawroot, and herbaceous food

production; (3) utility of prescribed fire in perpetuating a hard

mast species component in the forest; (4) creation or

perpetuation of escape cover through the use of prescribed

fire; (5) use of burned areas by bears; (6) behavior of bears
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 95

during prescribed burns; and (7) effective public education

programs to convey the benefits of prescribed fire for wildlife

habitat enhancement, forest resource production, hazard fuel

reduction, and public use.

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Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 97

Abstract

Fire is widely recognized as having been a major force

shaping the character of eastern hardwood forests.

Exclusion of periodic surface fires from mixed hardwood

stands for over 70 years has contributed to the gradual

succession of shade-tolerant species into the understory and

midstory of these stands. Following major disturbance to the

overstory, these shade-tolerant species or pioneer shadeintolerant species are able to out-compete oak regeneration

and dominate the next stand. Because of fire exclusion for

many decades, regeneration of oak-dominated stands has

been a major silvicultural problem. Recent research indicates

that a shelterwood-burn technique, which mimics the natural

disturbance regime that historically favored oaks, can be

used on productive upland sites to enhance the competitive

position of oaks in the advance regeneration pool. This

paper presents silvicultural guidelines for applying the

technique and discusses several options to meet the

management objectives of different landowners.

Introduction

Regenerating oaks on good quality sites has been a difficult

problem throughout the eastern United States for many

decades. Oak regeneration failures have generally been

attributed to either poor initial establishment of oak seedlings

or the slow juvenile growth of oak advance regeneration

when it is present (Abrams 1992, Lorimer 1993, Loftis and

McGee 1993). In the dense shade of mature mixedhardwood stands, oak seedlings and seedling sprouts do not

develop into competitive stems. Overstory removal by either

partial or complete cuttings releases well-established shade

tolerant regeneration and facilitates establishment of fastgrowing shade intolerant seedlings. Subsequent stand

development is to a mixed mesophytic forest with oak as a

minor component or altogether absent (McGee 1979;

Abrams 1992; Lorimer et al. 1994). This successional trend

is a relatively recent phenomena, developing in the past 75

years, and is tied to the exclusion of fire from eastern

hardwood forests (Little 1974; Van Lear and Johnson 1983;

Lorimer 1993).

Fire research in hardwoods has lagged far behind that in

pines, although several studies suggested that oaks were

ecologically adapted to frequent burning (Swan 1970,

Niering et al. 1970, Thor and Nichols 1974, Waldrop et al.

1987, Augspurger et al. 1987). All of these studies noted that

oaks resisted root kill by fire to a greater extent than their

competitors. Numerous authors, based on literature reviews,

accounts of early explorers and settlers, observed vegetative

patterns and responses, and other evidence, have

concluded that fire played an important role in the

development and maintenance of oak forests in the eastern

United States (Little 1974; Van Lear and Johnson 1983;

Crow 1988; Van Lear and Waldrop 1989; Williams 1989;

Abrams 1992).

In this paper, we will discuss the use of fire as a silvicultural

tool to regenerate oaks. Particular emphasis is placed on a

new regeneration method that utilizes partial harvesting

followed by prescribed fire.

Interactions Between Fire and Oaks

With the arrival of Indians in the eastern United States as

early as 12,000 years ago, fire became a more frequent

disturbance that shaped forest composition and structure

(Pyne 1982; Williams 1989). Indians used fire for many

reasons, including hunting, facilitating travel, stimulating

berry production, clearing land for agricultural crops, and as

a defense against enemies. The frequent, widespread use of

fire by Indians and the European settlers that followed them

created an environment favorable for the establishment and

maintenance of oaks. In the early 1900s, fire-suppression

efforts of the U.S. Forest Service and state forestry

commissions began to be successful in reducing the

frequency, extent, and influence of this powerful

environmental force. The forest changed dramatically as fire

was largely removed from the Eastern forest ecosystem.

Shade-tolerant and fire-intolerant species began to dominate

forest understories, overstory densities increased, and firesensitive species moved upslope from moist coves to xeric

slopes.

Frequent burning creates environments that favor oaks on

better-quality sites. Surface fires remove much of the midand understory strata in mature mixed hardwood stands,

reducing shading. Spring fires are especially effective

(Barnes and Van Lear 1998). Fire reduces the thickness of

the forest floor, preparing a favorable seedbed for caching of

acorns by squirrels and jays by (Darley-Hill and Johnson

1981, Galford et al. 1989). Fire reduces surface soil moisture

which discourages establishment of mesophytic species

(Barnes and Van Lear 1998) and may control insect

predators of acorns and new seedlings (Galford et al. 1989).

The presence of oaks encourage surface fires because of

the nature of their litter. An oak stand adds about 4.5 Mgha1

yr-1 of leaf litter to the forest floor (Loomis 1975). This litter

remains curly, creating a porous fuelbed for surface fires.

Unlike leaf litter of mesophytic species which forms a flat mat

upon compaction and decays rapidly, oak leaf litter

undergoes little decay during the winter. In regions where

snowpacks are heavy, oak litter recurls after snowmelt, once

Using Prescribed Fire to Regenerate Oaks

D.H. Van Lear 1, P.H. Brose2, and P.D. Keyser 3

1

Bowen Professor of Forestry, Department of Forest Resources,

Clemson University, Clemson, SC 29634-1003

2

Research Forester, USDA Forest Service, Warren, PA 16365

3

Wildlife Biologist, Westvaco Corporation, Rupert, WV 25984
98 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

again creating a porous fuelbed capable of carrying a

surface fire during the spring fire season (Lorimer 1989).

Because of the complexity of forest ecosystems, effects of

fire in hardwood stands vary. Fires in stands of mixed

composition have occasionally created oak-dominated

stands (Roth and Hepting 1943; Carvell and Maxey 1969;

Ward and Stephens 1989), probably because intense fires

controlled competition and stimulated rapid growth of oak

reproduction (Johnson et al. 1989; Lorimer 1989). In other

cases, species composition in young stands has been

altered very little by fire (Johnson 1974; McGee 1979;

Waldrop et al. 1985; Augspurger et al. 1987).

Effects of fire vary because of differences in season of

burning and fire intensity. Season of burning affects

physiological condition of the plant and the ability of species

to resprout. Hardwoods have the greatest ability to sprout

when carbohydrate storage in their roots is high, i.e., in the

dormant season. In the growing season, root reserves are

lower and sprouting vigor is less. Fire intensity is critical

because certain species, such as the oaks, can survive

higher intensity fires than their competitors (Brose and Van

Lear 1998) because their sprouts originate deeper in the soil

than those of their competitors (Hane 1999).

Using Fire to Encourage

Oak Regeneration

Understory Burning

Fire exclusion for most of the past century has altered stand

structure and composition of eastern hardwood stands.

Shade-tolerant and fire-intolerant species have gradually

encroached into the understory of oak-dominated stands.

Now, in the absence of periodic fire, there is no growing

space for oak reproduction, which may or may not be

present in the advanced regeneration pool. Van Lear and

Watt (1993) developed a theoretical silvicultural prescription

to encourage oak regeneration by burning the understory of

mature mixed hardwood stands near the end of the rotation.

Barnes and Van Lear (1998) tested this hypothesis in the

Piedmont of South Carolina and found that one burn early in

the growing season when leaves were expanding was as

effective as three winter fires in reducing density of

understory and midstory stems (fig. 1). The number of oak

rootstocks in the regeneration layer was increased by

burning, root/shoot ratios of oaks were enhanced, and

competitive woody species decreased. There was little

visible damage to boles of overstory oaks from the fires,

especially in the larger size classes.

Although oak regeneration was increased by understory

burning, it remained small and generally ephemeral. It

appeared that burning would have to be continued for a

relatively long period, i.e., perhaps 10 years or so, before

sufficient oaks of competitive size would be present in the

advance regeneration. Such an approach is handicapped by

the expense and risks of multiple prescribed fires, making it

a rather unattractive option for landowners and managers.

Nevertheless, if no advance regeneration exists in a stand,

periodic understory burning may provide a means to

establish oak seedlings and seedling sprouts, which could

then be encouraged to begin vigorous growth by the

following technique.

Shelterwood Cutting Followed

by Prescribed Fire

The Shelterwood-Burn Method

Oak-dominated stands on better quality sites in the southern

Piedmont and mountains often have abundant but small and

non-competitive oak reproduction in the regeneration layer.

When such stands are harvested by either clearcutting or the

shelterwood method, oak reproduction cannot compete with

Figure 1.—Reduction of midstory/

understory densities following three

winter burns and one spring burn in

mixed hardwood stands in the South

Carolina Piedmont.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 99

rapidly growing shade-intolerant species and/or wellestablished shade-tolerant species (Loftis 1983,

Abrams 1992, Schuler and Miller 1995).

A shelterwood-burn method (fig. 2) has recently been

developed in the Piedmont of the southeastern

United States to enhance the competitive position of

oak regeneration in such stands (Keyser et al. 1996;

Brose and Van Lear 1998; Brose et al. 1999a). This

method is based on the silvics and fire ecology of oak

and yellow-poplar regeneration and involves an initial

shelterwood harvest which removes roughly half of

the overstory basal area. All yellow-poplars are

removed in this first cut of the two-cut shelterwood

method, leaving the best dominant and codominant

oaks. Logging slash must be kept away from bases

of residual oaks by directional felling. This partial

harvest is followed by a 3- to 5-year waiting period,

after which a relatively hot growing-season

prescribed fire is run through the advance

regeneration.

Oak reproduction must be relatively vigorous and

free-to-grow to be competitive. We defined such oaks

as straight stems at least 1.3 m tall with no major

competitors within 3 m (Nix 1989). Oaks resist root

kill by fire better than yellow-poplar and other

competitors, especially as fire intensity increases (fig.

3) (Brose and Van Lear 1998). Density of free-togrow oaks exceeded 800 stems/ha and yellow-poplar

density was greatly reduced (up to 90%) in areas

burned in the spring with high intensity flames. In

contrast, winter burns provided little control of yellowpoplar and, even with a high intensity fire, oaks

density did not reach 300 stems/ha. Summer fire

resulted in substantial numbers of free-to-grow oaks

in all fire intensity levels, but especially in the two

medium intensity levels. High intensity summer fires

killed many of the smaller oak seedlings while low

intensity summer fires failed to control competition.

Oak reproduction will not be uniform over the entire

burned area. If free-to-grow oaks exceed 370/ha and

60% or more of the stocking plots have at least one

free-to-grow oak, the stand can be considered regenerated

with the likelihood that oaks will be a dominant component of

the next stand. One burn may not be enough if oak

dominance is desired in the new stand. Decades of fire

exclusion have allowed oak competitors to become firmly

established. If more oaks are desired, additional fires may be

prescribed as dictated by leaf litter accumulation. We believe

that oak dominance of the advance regeneration will

continually increase with repetitive spring burning at about 2-

year intervals.

We believe that a shelterwood cut, rather than a clearcut, is

the essential first step in this technique. The shelterwood

continues to produce oak litter which creates the continuous

fine fuel bed necessary to carry the subsequent fire. If a

clearcut had been used, foliage from yellow-poplar, a poor

medium for carrying surface fires, would dominate the forest

floor. In addition, shading from the shelterwood prevents

yellow-poplar regeneration from growing so large during the

interval before burning that it could not be killed by fire (Hane

1999).

In the 3 to 5- year period following the initial cut, logging

slash settles and loses it’s foliage, minimizing the risk of bole

damage to residual trees caused by flareups in heavy fuels

(Brose and Van Lear 1998; Brose et al. 1999a). Although

yellow-poplar was removed in the initial shelterwood cut, its

seed remains viable in the duff for up to 10 years (McCarthy

1933). Most of this stored seed germinates during the

waiting period and those seedlings are killed in the

subsequent fire. The waiting period also allows residual

overstory trees to recover from the shock of the initial cut

before they will be stressed again by burning.

Figure 2.—Schematic diagram of the shelterwood-burn technique. A =

high quality oaks; B = hickories, poor quality oaks, and yellow-poplars; C

= American beech, flowering dogwood, and red maple; D = mixed

hardwood regeneration dominated by yellow-poplar; and E = mixed

hardwood regeneration dominated by oaks.
100 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Advantages of the

Shelterwood-burn Technique

The shelterwood-burn method is attractive to landowners

because the initial cut of the shelterwood method produces

immediate income. A small portion of the profit is then used

to pay for the prescribed burn. Careful planning, e.g.,

consider using skid trails for fire breaks, before timber

harvest can reduce later prescribed fire costs. Removal of

the shelterwood after burning is at least as profitable as the

initial cut because the best oaks were retained and some

probably increased in value during the intervening years

before final harvest.

If the landowner’s goal is to manage for wildlife, the

shelterwood-burn method can be used to sustain hard mast

production while providing a source of palatable browse

during the regeneration period. Landowners can maintain

indefinitely the classic structure of the shelterwood while

stockpiling oak regeneration with periodic burns (Brose et al.

1999b). Many upland game and non-game species utilize

the mast, browse, and cover in a regenerating shelterwood

(Brose et al. 1999b; Lanham et al. 2000). A note of caution –

in areas where deer density is extremely high and early

successional habitat is relatively rare, the shelterwood-burn

method probably will not work because of over-browsing of

oak regeneration.

The shelterwood-burn system could be used to restore firemaintained ecosystems (Brose et al. 1999b). Frequent

growing season burns after the initial shelterwood cut would

gradually reduce woody regeneration (even the oaks) and

create a hardwood woodland or savannah, two habitats that

have become increasingly rare after decades of fire

exclusion (Buckner 1983; Pyne 1982; Van Lear and Waldrop

1989; Abrams 1992).

Research suggests that the shelterwood-burn technique may

be successful in other physiographic regions and on different

sites. Repeated burning in oak-pine communities on xeric

sites in the Cumberland Plateau reduced regeneration of red

maple and other non-oak species and promoted chestnut

oak regeneration (Arthur et al. 1998). On mesic sites in

Wisconsin, two spring burns reduced densities of sugar

maple and hophornbeam by 80%, while density of northern

red oak increased (Kruger 1992). Ward and Gluck (1999), in

Connecticut, observed that burning several years after a

shelterwood harvest favored oaks and reduced competition

from birch and shrubs. Hot fires in mountain laurel thickets in

the Northeastern United States that opened canopies, i.e., a

disturbance similar to a shelterwood harvest, allowed oak

reproduction to grow past the dense shrub layer (Moser et al.

1996).

Conclusions

Oak forests on good quality sites have been in decline in the

eastern United States for decades as more shade-tolerant

species gradually succeed oaks or shade-intolerant species

outgrew them after a major disturbance. Fire historically

played a major role in maintaining oak-dominated forests but

periodic surface fires have been excluded from these sites

for over 70 years.

Practical silvicultural prescriptions using fire for oak

regeneration have been lacking. Understory burning in

mature mixed hardwood stands creates environmental

conditions which should favor oak regeneration but

developing oak reproduction does not reach sufficient size to

be competitive when the overstory is removed.

A shelterwood-burn technique has recently been developed

which overcomes this major disadvantage. This technique

reduces the vigor of oaks’ competitors, especially if growing

season burns of relatively high intensity are used, and

provides conditions (increased light and reduced growth of

competitors) which allow oak regeneration to vigorously

resprout following the subsequent burn. It mimics a

Figure 3.—Mortality (%) of hickory, oak, red maple,

and yellow-poplar advance regeneration as fire

intensity increases within spring prescribed burns

conducted in shelterwood stands.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 101

disturbance pattern (i.e., partial overstory disturbance

followed by fire) that has shaped the composition of eastern

forests for millennia. The shelterwood-burn method is also

economically attractive because the initial shelterwood cut

yields an immediate income and the prescribed burn is low

cost compared to alternative treatments.

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Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 103

Abstract

Ridgetop pine communities of the Southern Appalachian

Mountains historically have been maintained by lightningand human-caused fires. With fire suppression for several

decades, characteristic stands are entering later seral

stages. They typically have an overstory of Table Mountain

(Pinus pungens) and/or pitch pine (P. rigida), a midstory of

chestnut oak (Quercus prinus), scarlet oak (Q. coccinea) and

blackgum (Nyssa sylvatica), and a shrub layer of dense

mountain laurel (Kalmia latifolia). Previous research

suggests that restoration of these communities can be

accomplished with high-intensity fires that open the forest

canopy and expose mineral soil. Three recent studies

examined plant-community response to high-intensity

prescribed fires. A series of corollary studies help to explain

some of the results of these field studies. High and mediumhigh intensity fires provided adequate sunlight for pine

seedlings, whereas medium-low and low intensity fires did

not. Post-burn duff was deep and did not vary by fire

intensity. We observed sufficient seedling densities to restore

pine-dominated stands after all but the highest intensity fires.

Many seedlings survived the first growing season as their

roots penetrated duff to reach mineral soil. Hardwood

rootstocks resprouted on sites treated with all fire intensities

and may out-compete pine seedlings for available resources.

High-intensity fires may have reduced mycorrhizal

abundance and moisture availability for new germinants.

Fires of lower intensity than previously recommended or

multiple fires of very low-intensity may best provide

conditions for pine regeneration, but additional research is

needed.

Introduction

Table Mountain pine (Pinus pungens) and pitch pine (P.

rigida) are ridgetop Appalachian endemics that historically

were maintained by lightning- and human-ignited fires.

Stands of Table Mountain and pitch pines present on the

landscape today were established by fires, logging, insects,

and diseases that occurred in the early twentieth century, the

most recent landscape-scale disturbances. Stand-replacing

fires have been nearly absent in the region since then

(Williams 1998). Sutherland and others (1995) found a Table

Mountain pine community in Virginia that was maintained by

major fires occurring approximately every 10 years until the

1940’s. Fire suppression and lack of other disturbance have

allowed ridgetop pine communities to succeed to hardwood

dominance and closed understories (Williams and Johnson

1992, Sutherland and others 1995, Turrill 1998, Williams

1998). As a result of these changes in dominance and

structure, Table Mountain/pitch pine woodlands are

recognized by the Southern Appalachian Assessment as one

of thirty-one rare communities in the Southern Appalachian

Mountains (SAMAB 1996).

Most research addressing the role of fire in Table Mountain

pine stands has been limited to post-wildfire studies, which

suggest that high-intensity prescribed fires that remove the

forest canopy and expose mineral soil will help ensure

successful regeneration. Zobel (1969) found that serotinous

cones opened in lightly burned areas, but that seedlings

survived only where fires killed overstory trees and erosion

exposed mineral soil. Likewise, Sanders (1992) observed the

greatest proportion of Table Mountain pine seedlings in highand moderate-intensity burn areas, where the canopy was

open and mineral soil exposed. Williams and Johnson (1992)

found that seeds were abundant on the ground in lightly

disturbed stands where no fire occurred. However, seedlings

were successful only on microsites with thin litter layers

(<1.5 in.) and where the canopy was more open than in

surrounding stands. Such microsites usually were created by

ice storms (Williams 1998).

Many ridgetop pine stands are located on National Forest

System land and National Parks, where prescribed burning

is possible and encouraged. Although many National Forest

and National Park land management plans prescribe highintensity fire for ridgetop pine communities, carrying out such

burns is difficult (Turrill 1998). Such prescriptions provide a

narrow window of opportunity and raise questions about

worker safety and smoke management (Waldrop and Brose

1999). In addition, some land managers avoid using highintensity fires because of the perceived risk of damaging

marketable hardwoods and the inability to control such fires

on steep slopes (Van Lear and Waldrop 1989). As a result,

high-intensity prescribed burning has had limited application

in the Southern Appalachian Mountains.

Williams (1998) suggested that Table Mountain/pitch pine

stands are in decline as a result of fire suppression policies

and inadequate understanding of the species regeneration

biology. To date, only three studies have conducted

prescribed burns to better understand the conditions

necessary for pine regeneration. Together, these three

studies examine community response to varying degrees of

fire intensity, as well as seedling establishment in varying

types of microhabitat. This paper examines the results of

Current Research on Restoring Ridgetop Pine Communities

With Stand Replacement Fire

Thomas A. Waldrop1, Nicole Turrill Welch2, Patrick H. Brose3, Katherine J. Elliott 4, Helen H. Mohr 1, Ellen A. Gray 5,

Frank H. Tainter 6

, and Lisa E. Ellis 6

1USDA Forest Service, Southern Research Station, Clemson, SC

2School of Public and Environmental Affairs, Indiana University,

Bloomington, IN

3

USDA Forest Service, Northeastern Research Station, Warren, PA

4

USDA Forest Service, Southern Research Station, Otto, NC

5

Department of Forestry, Wildlife and Fisheries, University of

Tennessee, Knoxville, TN

6

Clemson University, Department of Forest Resources, Clemson, SC
104 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

those studies to evaluate the fully open canopy and exposed

mineral soil conditions generally accepted as necessary to

regenerate ridgetop pines.

Current Research on Stand-replacement

Prescribed Burning

Effects of Fire Intensity on Table Mountain/

Pitch Pine Regeneration

Three studies have examined the response of Table

Mountain pine and pitch pine stands to stand-replacement

prescribed fire: Turrill (1998), Waldrop and Brose (1999), and

Elliott and others (1999). The burns conducted for these

studies varied in their effects on opening the forest canopy

and removing litter and duff. Comparisons of these field

studies allow an evaluation of the amount of pine

regeneration under natural conditions.

Several corollary studies provide insight to disturbance

history and methods of evaluating stands for their potential of

regeneration success. Waldrop and others (1999) conducted

a greenhouse study to evaluate the effects of shade and duff

on seedling establishment. Ongoing studies include the

dendrochronology of ridgetop pine stands across the

Southern Appalachians, seed biology of Table Mountain

Pine, and mycorrhizal associations in burned Table Mountain

pine stands. We will discuss the preliminary results of each.

The prescribed burn observed by Waldrop and Brose (1999)

was on the War Woman Wildlife Management Area of the

Tallulah Ranger District of the Chattahoochee National

Forest. Prior to burning, mean total basal area in study

stands was 123.0 ft2

 per ac. Hardwoods made up 98.8 ft2 of

this total and pines the remaining 38.8 ft2

. Chestnut oak was

the predominant hardwood and almost all pines were Table

Mountain pine. USDA Forest Service personnel conducted a

stand-replacement prescribed fire on a 850-ac unit in April

1997. The burn area included sharp ridgetops and steep

slopes with northeastern or southwestern aspects. The fire

was ignited by hand and by helicopter to create a ring fire

that reached greatest intensity within ridgetop Table

Mountain pine stands. The fire was large enough and its

intensity varied enough to allow comparisons of regeneration

success among areas burned at different intensities. This

ongoing study will include four additional burns: in Georgia,

South Carolina, and Tennessee during late winter 2000 or

the winter of 2000-2001.

Turrill (1998) studied a 7.5-ac prescribed fire on the

Grandfather Ranger District of the Pisgah National Forest.

The stand’s mean total basal area was 140.72

 ft per ac prior

to the burn. Hardwoods comprised 37.9 ft2

 per ac, and pines

102.8 ft2

 per ac. Blackgum was the predominant hardwood.

The pine component was 51 percent Table Mountain pine,

39 percent pitch pine, and 10 percent Virginia pine (P.

virginiana). USDA Forest Service crews used a combined

ring and head fire technique to burn the stand in May 1996.

Elliott and others (1999) studied a roughly 750-ac burn on

Wine Springs Creek on the Nantahala National Forest in

western North Carolina. The ridgetop community was

dominated by pitch pine, chestnut oak, scarlet oak, and red

maple, which comprised 49, 19, 8, and 8 percent of the total

basal area, respectively. This south-facing burn unit was

ignited by helicopter in April 1995 in a pattern to create a

mosaic of fire intensities. Low intensity fire occurred on the

lower slopes and flames reached into the crowns of trees

along the ridgetop.

Stands burned for Turrill (1998), Waldrop and Brose (1999),

and Elliott and others (1999) contained dense mountain

laurel shrub layers. Post-burn observations in both study

areas were completed in the first growing season after

burning.

The prescriptions applied in these studies produced four fire

intensities defined by Waldrop and Brose (1999): low,

medium-low, medium-high, and high. Waldrop and Brose

(1999) observed all four fire intensities, while Turrill (1998)

and Elliott and others (1999) observed only medium-low

intensity. Waldrop and Brose (1999) give a detailed

description of the discriminant functions used to classify fire

intensity, which generally can be described as follows:

Flames of low intensity fires never reached into the crown of

trees and uniformly burned the area. Medium-low-intensity

fires had flames slightly taller than those of low-intensity fire;

they burned less uniformly and produced hot spots where

flames reached into crowns and killed large trees. Flames of

medium-high intensity fires typically reached into the crowns

of all overstory trees. Flames of high-intensity fires generally

exceeded the crowns of overstory trees and carried from

crown to crown.

High- and medium-high-intensity fires reduced canopy cover,

leaving only 4.4 ft2 and 7.0 ft2

 per acre of basal area,

respectively (table 1). Mortality was high in all diameter size

classes following both high- and medium-high-intensity fires.

Sunlight reaching the forest floor may have been adequate

for seedling survival following fires of both intensities.

Medium-low- and low-intensity fires reduced canopy cover

(table 1), but residual basal area may be too high in all three

studies to allow stand replacement. Medium-low-intensity

fires reduced basal area to 48.3 ft2 per acre in Waldrop and

Brose (1999), 112.8 ft2

 per acre in Turrill (1998), and to 82.1

ft2 per acre in Elliott and others (1999). Low-intensity fires

had little effect on basal area, leaving 98.9 ft2

 per acre.

Mortality was greatest in lower d.b.h. classes (< 6 in. d.b.h.)

following fires of medium-low and low-intensity. Shade from

surviving trees may have prevented pine seedling survival.

Prolific hardwood sprouting was observed following fires of

all intensities (table 1). Waldrop and Brose (1999) reported

that all intensities top-killed the mountain laurel shrub layer;

Turrill (1998) and Elliott and others (1999) reported that

medium-low intensity fires did so. Generally, under all fire

intensities there were over 10,000 stems per acre, and they

were growing rapidly. Competition from these sprouts may

eliminate any pine regeneration after a fire of any fire

intensity. This result suggests that multiple, low-intensity fires
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 105

may be necessary to reduce hardwood abundance while

maintaining a seed source among large pines.

Post-burn counts of Table Mountain pine seedlings reported

by Waldrop and Brose (1999) and Turrill (1998) suggest that

fires were of sufficient intensity to open serotinous cones

throughout the burn unit, even in areas of low-intensity

burns. Post-burn pine density ranged from 1,400 to more

than 9,000 stems per acre (table 1). An unexpected result

was that the lowest pine densities were in areas burned at

the highest intensity. This suggests that cones were

consumed or seeds killed by intense heat, or that the

seedbed became less suitable.

Although plots in high-intensity burn areas had fewer

seedlings, if they are well dispersed, the 1,396 seedlings per

acre present in those areas should create pine-dominated

stands. However, Table Mountain pine seedlings were found

at only 51 percent of the sampling points, indicating that

portions of burned areas had no pine regeneration.

Hardwoods may dominate such areas. Plots in areas burned

at medium-high intensity also indicated low pine stocking (64

percent). If seedlings receive enough sunlight, pine density

and stocking levels in those areas burned at low and

medium-low intensities should be sufficient to create pinedominated stands.

Pitch pine does not have serotinous cones in the region

studied by Elliott and others (1999). Therefore, managers

were concerned that a winter or spring burn would consume

seeds already on the ground. In this study, over 1,100 pitch

pine seedlings per acre were present the year after burning,

indicating that the seed source was still viable. Among those

seedlings, however, fewer than 300 per acre survived. The

authors suggested that mortality was due to shading by the

surviving overstory, competition from sprouts, and a thick

duff layer that prevented seedling roots from reaching

mineral soil.

Pine seedlings regenerated on relatively thick duff following

all fire intensities in the Waldrop and Brose (1999) study.

Total litter and duff depth remaining after fires was 2.1, 1.5,

2.5, and 2.6 in. for the low-, medium-low-, medium-high-, and

high-intensity fires, respectively. The percentage of seedlings

with roots penetrating mineral soil was 71.1, 94.6, 63.0, and

56.1 for the same order of fire intensities (Waldrop and Brose

1999). Turrill (1998) observed pine regeneration on

approximately 3.6 in. of combined litter and duff. Waldrop

and Brose (1999) found that root systems of over 80 percent

of the sampled seedlings were able to penetrate duff up to

3.0 in., indicating that duff removal may not be as critical as

once thought. However, seedling survival was not tracked

beyond the first growing season in either study.

Corollary Studies

Dendrochronology

Little is known about the disturbance history of Table

Mountain/pitch pine stands. The species may have been

maintained by frequent low- to medium-intensity fires,

infrequent high-intensity stand-replacing fires, or a

combination of both. The dendrochronology study of

Sutherland and others (1995) provides valuable insight to fire

frequency and stand dynamics for one Virginia site. A similar

study is being done by Brose, Tainter, and Waldrop

Table 1.—Characteristics of Table Mountain and pitch pine stands one year following stand replacement

prescribed burning

 Fire Intensity Level

Medium Medium

Variable Low Low High High Source

Pine basal 25.7 26.1 4.8 0.0 Waldrop and Brose (1999)

area (ft2

/ac) 94.1 Turrill (1998)

42.1 Elliott and others (1999)

Hardwood 73.2 22.2 2.2 4.4 Waldrop and Brose (1999)

basal area 18.7 Turrill (1998)

(ft2

/ac) 40.0 Elliott and others (1999)

Total basal 98.9 48.3 7.0 4.4 Waldrop and Brose (1999)

area (ft2

/ac) 112.8 Turrill (1998)

82.1 Elliott and others (1999)

Hardwood 13,016 15,130 10,765 12,768 Waldrop and Brose (1999)

sprouts 929 Turrill (1998)

(num/ac) 12,354 Elliott and others (1999)

Pine seedlings 5,608 9,130 3,650 1,396 Waldrop and Brose (1999)

(num/ac) 3,117 Turrill (1998)

~300 Elliott and others (1999)
106 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

(unpublished study plan) in conjunction with the study sites

reported by Waldrop and Brose (1999). Cores have been

extracted from overstory trees and understory trees and

shrubs in two burn units in Georgia, two in South Carolina,

and one in Tennessee.

A preliminary analysis of stand dynamics in the two Georgia

units suggests a history of frequent disturbance that lasted

until the 1950’s (table 2). Pines in the dominant canopy

position are between 100 and 158 years old. However,

numerous smaller pines are between 50 and 100 years old.

Shrubs, particularly mountain laurel, are less than 50 years

old, and there are no pines younger than 50 years. The

frequency pattern of pine age classes (table 2) indicates that

pines were regenerating from the 1850’s through the 1950’s,

and that these stands were relatively open. Well-established

fire suppression policies in the 1950’s allowed the shrub

layer to become dominant and prevent continuing pine

regeneration. Restoration of these stands will likely require

some means to remove shrubs and competing hardwoods.

Seed biology

In the past, studies of prescribed burning assumed an

adequate seed source that did not vary among stands or

stand conditions. Any regeneration failures could have been

caused by an inadequate seed source. An ongoing study by

Gray, Rennie, and Waldrop (unpublished study plan) will help

identify stands that have an adequate seed source for

regeneration. Such studies will help managers determine the

abundance and viability of seed from a range of tree ages,

as well as from cones of different ages. Preliminary results

for seed viability are given in table 3. Seed viability was

moderate, generally between 20 and 50 percent, from cones

of all ages and from trees older than 10 years. Viability did

not appear to vary by age after trees reached 10 years.

However, viability seemed to increase as cones matured to 4

or 5 years old. These results indicate that, if cone numbers

are adequate, stands over a wide range of ages may be

considered as candidates for burning. A surprising result is

the presence of cones with viable seed on young trees.

Trees within the 5- to 10-year age class had 3-year-old

cones with 23 percent seed viability. This result suggests

that Table Mountain pines are adapted to regenerating under

regimes of low-intensity fires, which may occur every 5 to 10

years.

Mycorrhizae

The need for mycorrhizae is generally accepted for southern

pine seedlings grown in nurseries, but it has not been

studied for nontimber species such as Table Mountain pine.

Both ectomycorrhizae and vesicular-arbuscular mycorrhizae

may be necessary for survival of Table Mountain pine

seedlings, but their respective roles in Table Mountain pine

regeneration and their responses to high-intensity fires have

not been considered. Neary and others (2000) suggested

that fire intensity strongly affects the degree and duration of

reduced soil microbial activity. Neary and others (2000) also

suggested that after a period of low activity, microbial

populations increase in areas burned at high intensity to a

level much higher than in unburned areas, or in areas

burned at low intensity. This pattern may suggest that Table

Mountain pine seedlings will not develop for some time after

high-intensity burning; but that they may eventually benefit

from increased microbial populations. If so, prescribed burns

should be conducted far enough in advance of spring

germination to allow these populations to recover.

An ongoing study by Ellis, Tainter, and Waldrop (unpublished

study plan) is examining the relationship of fire intensity to

mycorrhizal development on Table Mountain pine roots.

First- and second-year seedlings were collected on 22 plots

established for the Waldrop and Brose (1999) study.

Seedling root biomass was quantified, ectomycorrhizal root

tips were characterized, and their presence compared by

seedling age, slope aspect, and microsite. Sites burned at

low and medium-low intensities had twice as many

mycorrhizal root tips (40 percent) than sites burned with

medium high and high intensities (22 percent). Two-year-old

seedlings had twice as many mycorrhizal root tips than oneyear-old seedlings. Seedlings on slopes of western aspect

had twice as many mycorrhizal root tips than those on

Table 2.—Age distribution of Table Mountain pines

sampled on two north Georgia sites

Age Class (yrs) n Pct of Sampled Trees

1-50 0 0

51-75 9 33

76-100 9 33

101-125 3 11

125-158 6 22

Table 3.—Percent viability of Table Mountain pine seed by tree age and cone age within a tree

 Cone Age All Cone

Tree age class (years) 2 years 3 years 4 years 5 years Ages

5 to 10 8 23 1 - -

11 to 25 20 32 41 23 27

26 to 50 33 11 24 56 31

51 to 75 29 20 34 36 30

75+ 29 13 54 39 33

All tree age classes 24 21 34 36
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 107

eastern aspects. Based on morphological and

histological examinations of mycorrhizal root tips,

Pisolithus tinctorius, Suillus granulatus, and

Cenococcum spp. were identified as the predominant

symbionts. Although these results are preliminary, they

may indicate that the very high-intensity fires had

adverse effects on mycorrhizal development, thus

reducing pine regeneration success.

Seedbed habitat

In order to assess seedling establishment, Waldrop

and others (1999) conducted a greenhouse study that

used shade and duff treatment combinations similar to

those observed in the field. Duff categories included

depths of 0, 2, and 4 in.; and shade levels included 0,

30, 63, and 85 percent shade. Table Mountain pine

seeds were collected on the Chattahoochee National

Forest in Georgia. Soil and duff were collected from a

recently burned Table Mountain pine stand on the

Sumter National Forest, South Carolina. Seeds were

germinated and allowed to grow under these

conditions for 3 months. We compared survival of

seedlings grown in the greenhouse to field seedling

survival in the burn described by Waldrop and Brose

(1999).

Figure 1 shows the total number of seedlings per plot

in all combinations of duff and shade at the end of the

90-day greenhouse study. Stem density typically was

greater in 2-in. duff than in bare soil or 4-in. duff. This

pattern remained constant for all except the 0-shade

category. In 0 shade, stem densities in pots with 2 in.

of duff were equal to stem densities in pots without

duff. Without shade, the mulching effect of a 2-in. duff

layer may not have been adequate to prevent

moisture deficit and seedling death.

Lack of shade reduced seed germination and the

survival of germinants, while heavy shade reduced

survival; more seedlings become established under

30-percent shade than under full light or the higher

shade levels. This pattern was constant among pots

with 2 and 4 in. of duff but differed among pots with no

duff (fig. 1). With no duff, fewer seedlings per pot

occurred under 30-percent shade than under no

shade, although this difference was not significant. Without

the mulching effect of duff, 30-percent shade may not be

adequate to prevent moisture deficit.

If germination and survival in the field follow the same

patterns as in the greenhouse, these data provide a partial

description of seedbed conditions necessary to establish

Table Mountain pine. Because of differences in study

designs, field results shown here do not provide a direct

comparison to greenhouse results. However, results of the

two studies are similar. In the field, stem numbers did not

vary significantly at different duff depths within a shade

category (fig. 2). Seedling numbers were not significantly

different between low- and medium-shade categories, but

both had significantly more stems than did the high-shade

category. Under high shade, stem density was less than

1,000 seedlings per acre at all duff depths, probably too few

to adequately regenerate a stand. Stem numbers in medium

and low shade ranged from 3,024 per acre for medium

shade with over 3 inches of duff to over 11,000 stems per

acre under medium shade and 1.6 to 3.0 inches of duff.

Each of these stem densities probably exceeds the minimum

needed to regenerate the stand.

The moderate levels of shade and duff that this study

suggests are optimum seedbed habitat differ somewhat from

previous recommendations. Although the exact fire regimes

necessary to create this type of habitat are unknown, these

results do not suggest that a single high-intensity fire is

Figure 1.—Seedling density per pot after the 90-day greenhouse

study for all combinations of shade level and duff depth.

Figure 2.—Seedling density in Georgia and South Carolina burn

units at the end of one growing season for all combinations of

shade and duff depth categories.
108 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

mandatory. Multiple lower-intensity fires could maintain an

overstory and seed source while reducing the duff without

exposing mineral soil.

Conclusions

The three studies described here represent the first attempts

to restore ridgetop pine communities in the Southern

Appalachians with prescribed stand replacement fires. Such

fires have been attractive for a number of reasons: they

provide a means of killing overstory trees and opening the

forest floor to direct sunlight; they provide the heat needed to

open serotinous cones; and they reduce thick duff layers or

expose mineral soil. However, none of the fires observed in

these studies should be considered successful for replacing

older stands of mixed pines and hardwoods with newly

regenerated stands of pines. Low-intensity fires observed by

Waldrop and Brose (1999) and medium-low intensity fires

observed in all three studies failed to kill more than a few

overstory trees. High intensity fires killed most overstory

trees but had few pine seedlings. Medium-high fires provided

abundant overstory mortality and pine regeneration.

However, fires of all intensities failed to control competition

from hardwood and shrub sprouts.

Competition and shading from hardwoods and shrubs that

sprout after burning may inhibit the development of a pinedominated stand. Post-fire sprouting occurred more

frequently in hardwood tree species (red maple, chestnut

oak, and scarlet oak) than in shrub species (mountain

laurel). The ability of Table Mountain and pitch pine

seedlings to compete with the regeneration of other species

is unknown. Frequent burning may be necessary to reduce

hardwood sprout vigor.

The corollary studies presented here provide indirect

evidence that ridgetop pine communities may be restored by

frequent burning. The dendrochronology study shows that

pines in study stands were uneven-aged and had

regenerated frequently until the time of fire exclusion. The

seed biology study suggests that a viable seed source is

present over a wide range of tree ages and in cones that

have been on trees for up to 5 years. This study also shows

that very young trees produce viable seed, suggesting an

adaptation to frequent burning. Studies of seedbed habitat

and mycorrhizal populations provide evidence that the

severe conditions produced by high-intensity burning are not

necessary and may be detrimental to regeneration. Moisture

may be limited due to lack of mycorrhizal tips on roots, loss

of a mulching effect from the duff, and direct sunlight

reaching the forest floor. Moderate levels of shade with some

duff present were optimum for seedling survival, both in the

greenhouse and in the field. These conditions may have

been common in pre-1950’s stands that burned often. Thick

duff was a barrier to seedling development in all field studies

but seedling roots can penetrate duff up to 3 inches thick.

The results presented here suggest that ridgetop pine stands

were created by lower-intensity fires than once were thought

necessary, and that such fires would aid in stand restoration.

Low-intensity prescribed fires, which can be used when the

lower layers of the forest floor are moist, are less dangerous

and present a larger window of opportunity than highintensity fires. Low-intensity fires also decrease erosion

potential on steep slopes and loss of site productivity.

There is still much to learn about restoring ridgetop pine

stands. If seedlings continue to survive among sprout

competition, a single medium-high intensity fire may prove

sufficient. However, these results were drawn from studies

which tracked seedling survival and overstory mortality for

only one growing season. More research is necessary before

definitive fire plans can be developed for ridgetop pine

communities. These studies should apply prescriptions to

achieve medium-high intensity burns and observe post-burn

canopy cover, seedling density, seedling rooting depth, and

seedling survival over several growing seasons. Additional

research also is needed to test fires in other seasons and

multiple low-intensity burns. Such fires should be conducted

so as to control hardwood and shrub sprouting while

maintaining healthy overstory pines as a seed source.

Serotinous cones can open with the low-intensity fires (Zobel

1969, Waldrop and Brose 1999), indicating that the reduced

fuel loads after multiple burns would not reduce fire intensity

too much to allow cones to open.

Many questions remain about the ecology of Table Mountain

and pitch pines. In particular, the competitive ability of these

species is unknown. Physical, chemical, and biological

properties of soils in ridgetop stands are likely to be affected

by regeneration burns. These properties may affect seedbed

conditions but they have not been studied. Finally, natural

disturbances, other than fire, may have played an historical

role in perpetuating the species; so alternative management

strategies may also improve regeneration.

Acknowledgment

Partial funding for the research described here was awarded

by the Department of Interior/Department of Agriculture,

Forest Service Interagency Joint Fire Science Program.

Literature Cited

Elliott, Katherine J.; Hendrick, Ronald L.; Major, Amy E.;

Vose, James M.; Swank, Wayne T. 1999. Vegetation

dynamics after a prescribed fire in the southern

Appalachians. Forest Ecology and Management. 114:

119-213.

Neary, D.G.; DeBano, L.F.; Ffolliott, P.F. 2000. Fire impacts

on forest soils: a comparison to mechanical and

chemical site preparation. In: Moser, W. Keith., ed.

Proceedings of the 21st Tall Timbers Fire Ecology

Conference; 1998 April 14-16; Tallahassee, FL. Tall

Timbers Research, Inc. (In press).

Sanders, Greg L. 1992. The role of fire in the regeneration

of Table Mountain Pine in the Southern Appalachian

Mountains. Knoxville, TN: The University of Tennessee,

Knoxville. 123 p. Thesis.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 109

Southern Appalachian Man and the Biosphere (SAMAB).

1996. The Southern Appalachian Assessment

Terrestrial Technical Report. Report 5 of 5. Atlanta:

USDA Forest Service, Southern Region.

Sutherland, E.K.; Grissino-Mayer, H.; Woodhouse, G.A.;

Covington., W.W.; Horn, S.; Huckaby, L.; Kerr, R.; Kush,

J.; Moore, M.; and Plumb, T. 1995. Two centuries of fire

in a southwestern Virginia Pinus pungens

community. In: Inventory and management techniques in

the context of catastrophic events: altered states of the

forest. Hypertext Proceedings, University Park,

Pennsylvania State University, Center for Statistical

Ecology and Environmental Statistics.

Turrill, Nicole L. 1998. Using prescribed fire to regenerate

Pinus echinata, P. pungens, and P. rigida

communities in the Southern Appalachian Mountains.

Knoxville, TN: The University of Tennessee, Knoxville.

207 p. Dissertation.

Van Lear, David H.; Waldrop, Thomas A. 1989. History, use,

and effects of fire in the Southern Appalachians. Gen.

Tech. Rep. SE-54. Asheville, NC: U.S. Department of

Agriculture, Forest Service, Southeastern Forest

Experiment Station. 20 p.

Waldrop, Thomas A.; Brose, Patrick H. 1999. A comparison

of fire intensity levels for stand replacement of Table

Mountain pine (Pinus pungens Lamb.). Forest Ecology

and Management. 113: 115-166.

Waldrop, Thomas A.; Mohr, Helen H.; Brose, Patrick H.;

Baker, Richard B. 1999. Seedbed requirements for

regenerating Table Mountain pine with prescribed

fire. In: Haywood, James D., ed. Proceedings of the tenth

biennial southern silvicultural research conference; 1999

February 16-18; Shreveport, LA. Gen. Tech. Rep. SRS30. Asheville, NC: U.S. Department of Agriculture, Forest

Service, Southern Research Station: 369-373.

Williams, C.E. 1998. History and status of Table Mountain

pine-pitch pine forests of the Southern Appalachian

mountains (USA). Natural Areas Journal. 18: 81-90.

Williams, Charles E.; Johnson, W. Carter. 1992. Factors

affecting recruitment of Pinus pungens in the

southern Appalachian Mountains. Canadian Journal of

Forest Research. 22: 878-887.

Zobel, D.B. 1969. Factors affecting the distribution of

Pinus pungens, an Appalachian endemic. Ecological

Monographs. 39: 303-333.
110 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

The Kentucky State Nature Preserves Commission is

charged with the responsibility to inventory the state for rare

species and unique ecological communities and to protect

them in perpetuity in a system of nature preserves.

Presently, there are thirty-nine state nature preserves and

nine of them protect barrens communities. Barrens can be

described as sparsely wooded ecological communities with

thin rocky soils underlain by sandstone or limestone. Barrens

are characterized by tree cover ranging from 25 to 60

percent and a ground cover dominated by perennial grasses

and prairie forbs, some of which are listed as rare species in

Kentucky. Protecting these preserves in perpetuity requires

active management to maintain open habitat which sustains

the grasses and forbs. Years of fire suppression have

resulted in canopy closure and the disappearance of some

species. Efforts in Kentucky are focused on restoration of

overgrown barrens. Prescribed fire is one tool to reduce

canopy and midstory cover. Fire also removes litter that can

inhibit seedling establishment and limit plant growth.

Examples of barrens communities in Kentucky will be

presented along with agency goals for restoration and

management.

Barrens Management and Restoration: A Kentucky Example

Joyce Bender 1

1

Kentucky State Nature Preserves Commission, 801 Schenkel

Lane, Frankfort, KY 40601, ph.: 502-573-2886, email:

Joyce.Bender@mail.state.ky.us
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 111

Abstract

Fires wound trees; but not all of them, and not always.

Specific fire behavior and differences among tree species

and individual trees produce variable patterns of wounding

and wound response. Our work focuses on the relationships

between fire behavior and tree biology to better understand

how hardwood trees resist injury to the lower stem and either

survive or succumb to low-intensity fire. Our objectives here

were to 1) define and describe the wounding process, 2) to

describe compartmentalization and wound closure and 3) to

discuss species-specific differences among several common

hardwood trees in the resistance to injury and resilience after

wounding. Characteristics of fire scars are summarized.

Introduction

Prescribed fire is a frequently discussed and under-utilized

tool to restore upland oak (Quercus) communities in Central

Hardwoods forests. One barrier to the use of prescribed fire

is a lack of understanding of how fire, especially low-intensity

prescribed fire, affects individual trees and tree species.

Fires wound tree stems (Gustafson 1946, Kaufert 1933,

Nelson and others 1933, Wendel and Smith 1986), but not

all of them, and not always (Smith and Sutherland 1999).

Patchiness of fire behavior and different susceptibilities

among individual trees produce variable patterns of

wounding. Additionally, tree species differ in their

effectiveness to resist injury and the spread of infection after

wounding. Knowledge about potential wounding from fire

may be an important consideration in forest managers’

decision-making processes. For example, some managers

may be concerned about loss of economic value through

damage to potential high-quality veneer logs. Alternatively,

those burning to restore the structure and composition of oak

communities undergoing succession to mesic forest types

might want to maximize injury to undesirable species (such

as maple (Acer)). Here, we define and describe the

processes of fire-caused wounding, the generalized

response of hardwood trees to wounding, and speciesspecific differences in protection from fire injury and

effectiveness in the tree wound response.

How Fires Wound Trees

In hardwood forests, fires typically burn in light fuels

(hardwood litter) and are low in intensity (Komarek 1974).

Not all trees are wounded by any one low-intensity fire

(Smith and Sutherland 1999) because within a fire perimeter

there are usually unburned patches; obviously, trees in these

patches are unlikely to be injured. The probability that a

hardwood tree will be wounded during a fire depends on

many factors— primarily weather, fire behavior and bark

characteristics. Fire wounds the vascular cambium by lethal

heating, which depends on flame length, fire temperature,

and duration of heating (Ryan 1998). These fire behaviors

are in turn affected by fuel quantity and moisture content,

ambient temperature, wind, slope, and in prescribed fires, by

ignition pattern (Ryan 1998). Wildfires differ from prescribed

fires. By definition and by regulation, prescribed fires should

only be ignited when they are judged controllable, under

specific parameters of weather and fuel conditions. Thus, the

fire manager igniting a prescribed fire has some control over

fire behavior and by extension, the probability of tree injury

and damage.

By itself, scorch is not an indicator of injury. Fire injury is the

result of fire causing a wound, a disruption of living tissues

including the vascular cambium that results in an impairment

or loss of function. These wounds are typically referred to as

fire scars. However, not all scorched trees are injured;

scorch is simply a sign or indicator of a fire occurrence.

Further, a scorched or even an injured tree is not necessarily

damaged: damage involves a loss of desirable wood

characteristics, value, or usefulness. The determination of

damage is relative to management goals. Damage due to a

fire scar may be essentially zero to a tree in a conservation/

preservation area, be worth thousands of dollars to an

identical tree managed to produce high-quality veneer, or

actually be desirable for wildlife management goals (e.g., to

promote habitat for animals that dwell or feed under dead

bark or in cavities). For timber management, methods to

estimate cull as a result of fire wounding were developed by

Hepting (1941) and Loomis (1973,1974). Notably, estimated

cull is virtually negligible when externally visible wounds are

less than 2 inches inches width at 1 foot aboveground

(Hepting 1941). Loomis (1974) suggested that oak trees with

wounds less than 6 in wide are unlikely to lose quality and no

more than 3 board feet in volume, and that pole-sized trees

were unlikely to lose any quality. Unpublished data of the

authors indicates that most prescribed fire-caused wounds

were less than 1 inch in width.

Most fire-caused injuries from low-intensity fires result from

heating without combustion. Smith and Sutherland (1999)

found that most injury to stems in Quercus sp. from two

prescribed fires resulted from heat conducted through bark,

and not flaming combustion. Heat transfer properties govern

resistance to fire-caused injury (Ryan 1998). The single most

important biological factor determining resistance to fire

injury to stems is bark thickness (Hare 1965, Jackson and

others 1999, Ryan 1998, Ryan and Reinhardt 1988, Vines

1968). Bark serves as an insulator that slows heat

Resistance is not futile: The response of hardwoods

to fire-caused wounding

Elaine Kennedy Sutherland1 and Kevin T. Smith2

1

Project Leader, USDA Forest Service, Rocky Mountain

Research Station, Forestry Sciences Laboratory, PO Box 8089,

Missoula, MT 59807

2

Plant Physiologist, USDA Forest Service, Northeastern

Research Station, Forestry Sciences Laboratory, 271 Mast Rd.,

Durham, NH 03824.
112 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

conduction to the vascular cambium (Spalt and Reifsynder

1962). The thicker the bark, the longer the heating period

required to injure underlying cambium (Uhl and Kaufman

1990, Vines 1968).

In general, bark thickens as trees grow in diameter, and

resistance to fire-caused wounding increases (Harmon 1984,

Hengst and Dawson 1994, Pausas 1997, Spalt and

Reifsynder 1962). Bark thickness is dependent on diameter,

not age (Hengst and Dawson 1994); suppressed trees the

same age as dominant trees have relatively thinner bark and

hence greater susceptibility to fire injury (Harmon 1984).

Consequently, ability to resist fire injury has been attributed

to stem diameter and to variability among tree species (Hare

1965, Hengst and Dawson 1994, Pausas 1997, Spalt and

Reifsynder 1962).

Some tree species fundamentally have thicker bark than

others, and as such have greater resistance to fire injury.

Because texture is so variable thickness is difficult to

quantify, but general patterns among species have been

noted (e.g., Harmon 1984, Hengst and Dawson 1994). For

example, upland species generally have thicker bark than

bottomland species, perhaps because fire is typically more

frequent at upland sites compared to bottomland sites

(Hengst and Dawson 1994, Jackson and others 1999).

Comparisons of data presented by Harmon (1984), Hengst

and Dawson (1994), Jackson and others (1999), and Spalt

and Reifsynder (1962) reveal certain patterns. As a group,

white oaks (Quercus subgenus Lepidobalanus) have the

thickest barks of the central hardwoods species, followed by

the red oaks (Quercus subgenus Erythrobalanus). Examples

of thinner barked species include American beech (Fagus

grandifolia), maple species (Acer), hickory species (Carya),

flowering dogwood (Cornus florida), and black cherry

(Prunus serotina).

Additionally, the rate of bark thickening on the lower stem

varies among species and primarily determines resistance to

heat-caused injury; the sooner bark thickens at the base, the

earlier in life the stem gains resistance (Hare 1965, Harmon

1984, Hengst and Dawson 1994). For example Hengst and

Dawson (1994) showed that tulip poplar (Liriodendron

tulipifera) has very thin bark when small but has a very rapid

rate of bark thickening. Thus, seedling and sapling-sized

tulip poplar trees are susceptible to topkilling by fire (Barnes

and Van Lear 1998, Brose and Van Lear 1998), but treesized tulip poplar has been described as particularly resistant

to fire-caused wounding (Gustafson 1946, McCarthy 1933,

Nelson and others 1933). By contrast, silver maple (Acer

saccharinum) also has very thin bark when small but a slow

rate of bark thickening, which results in thin bark throughout

its life (Hengst and Dawson 1994); silver maple is highly

susceptible to fire injury (Auclair and others 1973).

The distribution of bark along and around the stem is also

variable among species. Bark texture can vary from smooth

to fissured and can vary significantly within as well as among

species (Howard 1977). For trees with deeply fissured bark,

such as chestnut oak (Q. prinus), the cambium directly

beneath the fissures is more susceptible to injury, and

multiple, discontinuous fire scars may occur around the stem

(Smith and Sutherland 1999, Stickel and Marco 1936).

Factors that contribute to variability in bark texture include

tree vigor and growth rate, age, and height on the tree

(Howard 1977). In regimes characterized by low-intensity

surface fires, thick bark at tree bases provides protection

from heating and a competitive advantage over trees with

thin bark at the base, but since fires typically have low

flames heights, bark rapidly thins with increasing height

(Harmon 1984, Jackson and others 1999). For example, bur

oak (Quercus macrocarpa) has relatively thick bark near the

ground (but thinning with height); it grows in oak savannas

that are maintained by frequent fire. By contrast, water oak

(Quercus nigra) has relatively thin bark near the ground, and

typically grows in bottomland forests where fire rarely occurs

(Jackson and others 1999).

Heat conduction properties of bark tissue play a role in

resistance to heating and injury of the vascular cambium

during fires, and these properties vary among species (Hare

1965, Harmon 1984, Hengst and Dawson 1994). For

example, for a given bark thickness, red maple (Acer

rubrum) conducts heat relatively quickly (Hare 1965). Heat

conduction properties have been expressed as thermal

conductivity (the ability of a material to transfer heat),

specific heat (the ability to absorb heat), and thermal

diffusivity (ratio of thermal conductivity to the product of

specific heat and bark density) (Martin 1963, Harmon 1984).

The ratio of outer bark (rhytidome) to inner bark (phloem)

increases with tree diameter in many species, and this factor

affects heat conduction properties: rhytidome conducts heat

more slowly (Hare 1965). Thus, trees with substantial

rhytidome such as chestnut oak (Quercus prinus) are more

heat-resistant than trees with little rhytidome such as

American beech (Fagus grandifolia) (Stickel 1936). However,

bark thickness alone remains the simplest, best predictor of

heat conduction (Vines 1968).

How Trees Respond to Fire Injury

Tree survival after mechanical injury, including wounding

caused by fire, depends on the internal boundary-setting

process known as compartmentalization (Shigo 1984).

Compartmentalization in the wood of living trees resists the

spread of decay and loss of normal wood function, and

minimizes the extent of injury. Tree species vary in ability to

compartmentalize injury (Shigo 1984, Shortle and others

1996) and hence in resilience to fire-caused injury. For

example, trees in the white oak group (Quercus subgenus

Lepidobalanus) are unusually effective at

compartmentalizing decay, which places them at a

competitive advantage over other injured trees (Abrams

1996). Understanding the role of compartmentalization

effectiveness following fire may be critical to understanding

patterns of tree survival and changes in wood quality

following fire.

Compartmentalization is a set of processes that integrate

tree anatomy, physics, and inducible changes in tree

physiology. These processes may be divided into those that

occur in wood present at the time of wounding and those that
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 113

occur in wood formed after wounding. Tree survival depends

on maintaining a healthy vascular cambium, the thin layer of

generative cells located beneath the bark. As the cells of the

vascular cambium divide, wood (xylem) is formed to the

inside of the cambium and inner bark (phloem) is formed to

the outside. When part of the vascular cambium and

associated tissues are overheated and killed by fire, the first

changes are physical and immediate: water columns

(normally held under tension in functional wood vessels)

snap which is dangerous for the tree because drying kills

living sapwood cells and provides a favorable environment

for the spread of microbial infection. Because of cellular

architecture and inducible changes in wood physiology,

these abrupt changes in water tension induce the immediate

formation of plugs and the eventual formation of chemical

boundaries that resist both wood drying and the spread of

microorganisms that lead to the breakdown of wood. Some

of the physiological changes involve the oxidation and

polymerization of phenolic compounds that then discolor or

stain the wood (Smith 1997).

After wounding, probably in response to changes in plant

growth regulators (Smith and Shortle 1990), the surviving

vascular cambium produces an anatomically distinct barrier

zone (Shigo 1984) that can appear as a “false ring” in the

wood. This barrier is most effective in resisting the outward

loss of wood function and the spread of microorganisms.

Effective barrier zones limit wood discoloration and decay to

wood present at the time of injury. Compartmentalization

allows for the continued survival of the vascular cambium

and the generation of new wood. These boundaries,

although frequently effective at resisting the spread of

infection, can and do fail. The effectiveness at

compartmentalization depends on the severity of the wound,

the tree species involved, and on the individual tree.

For years following the injury, conspicuously wide growth

rings may be produced at the wound margins, apparently to

both speed closure of the wound and to provide additional

mechanical support for the wounded stem. Wound closure,

although apparently not essential for tree survival, aids tree

functioning by restoring the continuity of the vascular

cambium and wood formation around the tree perimeter.

Closure also seals off the wound and infected wood from the

outside atmosphere, reducing the partial pressure of oxygen

and increasing the partial pressure of carbon dioxide. These

changes in the internal atmosphere tend to reduce the rate

of the anaerobic wood decay process. Research on northern

hardwoods indicates compartmentalization is more effective

in sugar and red maple and red oak than in paper and yellow

birch, following equivalent levels of storm injury (Smith,

unpublished data).

Characteristics of Fire Scars

In earlier work (Smith and Sutherland 1999), we

characterized fire scars in small, dissected oaks in southern

Ohio that had been exposed twice to prescribed fire. We

learned that: 1) the prescribed low-intensity fires wounded

some trees, but because heat was unevenly distributed, not

all trees were scorched, and not all scorched trees were

wounded; 2) because flame heights were low, most wounds

were small and low to the ground (about 1 foot in height

above groundline); 3) wounds resulted in typical defense

processes against infection, including compartmentalization

and closure; and 4) nearly all wounds were caused by

heating through the bark, which often persisted over the

heat-killed tissue after wounding, sometimes for years, thus,

small wounds that rapidly compartmentalize may only be

apparent when the tree is felled and dissected.

Implications

The resistance of any hardwood tree stem to fire-caused

wounding is primarily determined by bark thickness.

Generally, as stems increase in diameter, bark thickness

concomitantly increases along with resistance to fire-caused

wounding. Additionally, there are species-specific differences

in bark thickness. Thus, there is a gradient of susceptibility:

small, thin-barked trees are most susceptible to wounding

and large, thick-barked trees are least susceptible.

Many researchers have suggested that fire is an important

process in promoting oak dominance in upland oak types, for

example Abrams (1992) and Lorimer (1985, 1993).

Differences among species in resistance to wounding by fire

and the effectiveness of wound response (resilience to

wounding) is probably an important determinant of species

composition and dominance in forests exposed to fire.

Jackson and others (1999) provided evidence for this idea.

They pointed out that for any tree, the greatest improvement

in survival probability occurs at sapling sizes, and then

demonstrated that the rate of overall growth and bark

thickening is more rapid in oak species typical of fire-prone

habitats than those associated with moist habitats (e.g.,

bottomlands) that burn infrequently. They showed that

resistance is higher in species typical of fire-prone habitats,

and suggested that other defenses would probably be

quantifiable. We believe that resilience to injury (effective

compartmentalization) is one of those defenses, and would

likely follow similar patterns of greater resilience in species

typical to fire-prone habitats compared to those where fire is

rare.

Conclusions

Any fire, whether wild or prescribed, can injure a tree, but not

all trees are wounded by fire. Fires in hardwood forests are

typically low in intensity and patchy in distribution. Even

when scorched, not all trees are wounded. Bark is an

insulator from heat and if scorched, the underlying cambium

may not be damaged. Most wounds, when they occur, are

near the ground, small in size, and rapidly

compartmentalized: few are externally visible. In addition, not

all wounds are caused by fire: other events also

mechanically wound trees, including impacts from treefalls

(e.g., windthrows) and management activities, including

logging. Because damage is relative to management

objectives, not all injuries result in damage. In the

development of fire prescriptions and in implementing fires,

managers have a degree of control over potential for firecaused injury based on their desired outcome.
114 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Better understanding of species-specific resistance and

resilience to fire injury are essential to improving predictions

about the effect of prescribed fire on surviving trees. To this

end, the authors are currently conducting research on the

characteristics of prescribed fire-caused wounds, resistance

to injury, and comparative effectiveness of

compartmentalization in central hardwood species.

Acknowledgments

This research was funded by Ecosystem Management

monies provided by the USDA Forest Service to the

Northeastern Research Station’s Ecosystem Management

project “Restoration of mixed-oak forests in southern Ohio

with prescribed fire.”

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116 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Abstract

Previous to 1996, wildland fires were effectively excluded

from Great Smoky Mountains National Park (GSMNP) for

over 60 years. The National Park Service now recognizes

the importance of fire as a natural disturbance and considers

fire a critical disturbance factor in GSMNP. A Management

Ignited Prescribed Fire Program, part of the Park’s Fire

Management Plan, was created to address some resource

management objectives. Prescribed fire is now being used

as a tool to enhance natural resources such as rare plant

populations. Four rare plant species have been managed by

prescribed fire since 1997 and all populations have

responded with increased plant numbers and vigor. Based

upon our success, two additional species are scheduled for

management by prescribed fire in 2000.

Introduction

Fire has been excluded from Great Smoky Mountains

National Park (GSMNP) since the Park’s establishment

(1934). An approved Fire Management Plan (1996), and the

institution of a Fire Management Program, has allowed

GSMNP to restore an important natural disturbance to various

natural communities as well as individual plant populations. A

Rare Plant Program was initiated in 1989 to monitor rare

plant populations and, if these species were observed to be

declining, to manage these populations in order to stabilize

or reverse the trend. Currently, the following four rare plant

species are being managed with prescribed fire:

• Daisy-leaved moonwort (Botrychium matricariifolium A.

 Braun)

• Bittercress (Cardamine flagellifera O. E. Schulz)

• Goldenseal (Hydrastis canandensis L.)

• Indian grass (Sorgastrum nutans (L.) Nash)

Southern Appalachian cove forests are typically affected by

small-scale, mild disturbances such as tree falls, and many

rare plant species require some level of disturbance in order

to survive (Denslow 1987). Through our monitoring efforts,

we determined that two species – Daisy-leaved moonwort

and Bittercress – appeared to require some leaf litter

disturbance to stimulate germination and thrive. We chose

prescribed fire to create disturbance to the leaf litter although

fires were relatively infrequent within these mountains in

general, and north-facing lower slopes have the lowest

incidence of fire (Harmon 1982). Three populations (Daisyleaved moonwort, Bittercress, and Goldenseal) are located

on north-facing slopes (elevation range 381-686 meters) in

mixed mesophytic hardwoods. In addition, GSMNP’s single

population of Goldenseal was infected with a native fungus,

Streptobotrys streptothrix, and fire was used to reduce the

sclerotia of the fungus which overwinter in leaf litter. The

fourth species, Indian grass, has been negatively affected by

annual mowing and competition from non-native Fescue

(Festuca pratensis). Prescribed fire was used to stimulate

flowering in the native grass.

Methods

Daisy-leaved moonwort

Two populations of Daisy-leaved moonwort (Botrychium

matricariifolium A. Braun), a small, primitive fern (3-19 cm in

height), are known to occur in GSMNP. This species is listed

as significantly rare in North Carolina and as a species of

special concern in Tennessee. One population is comprised

of 42 plants and the other approximately 114 plants, or

fewer, depending on the year.

Their habitat is second-growth tulip poplar (Lirodendron

tulipifera), red maple (Acer rubrum), and Eastern hemlock

(Tsuga canadensis). We chose the larger population for

management. A trail bisects the population creating a light

gap. Disturbance to the leaf litter layer periodically occurs

from episodes of heavy rainfall, wild boar rooting (usually in

the winter), and hikers. The population is north-facing with a

slope of 5-50 percent (elevation 686 meters).

Two permanent transects (21 meters and 10 meters) were

established in the study population in 1991, and plants

occurring within three meters of each transect were mapped.

Since 1991, plants were most abundant in locations devoid

of leaf litter and duff. Due to a severe decline in plant

numbers in 1997 (67 percent), fire and raking were selected

as management tools. Mapped plants were used as a guide

to establish three 7 meter x 10 meter treatment plots to

which we applied three different management techniques:

(1) fire to decrease leaf litter and duff, (2) raking, and (3) no

treatment in March 1998.

Bittercress

Bittercress (Cardamine flagellifera O. E. Schulz), a

stoloniferous perennial, is listed as state threatened in

Tennessee. The Park has three known populations and two

are actively monitored due to potential threats (e.g. wild boar

rooting and deer browsing). Baseline data were collected at

one population in 1989 and censused in 1991.

The study population is on a north-facing slope in secondgrowth mesic hardwoods comprised of tulip poplar, red oak

(Quercus rubra) and white oak (Quercus alba) (elevation 610

meters). Due to a decrease in rosettes observed in April

1997 (16 percent), a portion of the population was burned in

September 1997. The study population was re-censused in

April 1998.

Managing Rare Plant Populations With Fire

in Great Smoky Mountains National Park

Janet Rock 1

1Botanist, Great Smoky Mountains National Park, 107 Park

Headquarters Road, Gatlinburg, TN 37738
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 117

Goldenseal

Goldenseal (Hydrastis canadensis L.) is a highly valued

perennial herb with thick yellow rhizomes. This plant is listed

in the Convention on International Trade of Endangered

Species (CITES) Appendix II (USFWS/OSA 1997), and is

state-listed as endangered in North Carolina and threatened

in Tennessee due to over-collection and habitat loss. The

Park has one population comprised of two colonies

(subpopulations 1 and 2). Goldenseal prefers rich moist

woods, and in GSMNP it is found growing in mesic secondgrowth woods comprised of sugar maple (Acer saccharum),

American beech (Fagus grandifolia), red oak (Quercus

rubra), and Eastern hemlock on a north-facing slope of 50

percent (elevation is 381 meters).

Data has been collected each year since 1989. In 1991 a

native fungal disease, identified as Streptobotrys streptothrix,

was observed on the stems and leaves throughout the

population (Alan Windham, University of Tennessee

Agricultural Extension Agency). Subpopulation 1 showed the

highest incidence of infection, and by June 1991 it was

observed that the total number of stems had decreased by

30 percent in just one year.

Based on Windham’s recommendation, in September 1991

leaves and stems were clipped, bagged, and removed from

the site in both subpopulations. Leaf litter directly under the

stems was bagged and removed from the site as well. In

addition, in March 1998, a low-intensity fire was set at

subpopulation 1 in order to consume sclerotia of the fungus

overwintering in the leaf litter layer.

Indian grass

Indian grass (Sorgastrum nutans (L.) Nash) is a warmseason grass. Increase in Indian grass stems after burning is

well-documented (Dix and Butler 1954; Robocker and Miller

1955; Kucera and Ehrenreich 1962; Hadley and Kieckhefer

1963; Anderson and others 1970). Scattered populations

occur on the western end of the Park, particularly in open

fields in Cades Cove. It has been suppressed as a result of

haying operations since the 1940s, and is out-competed by a

non-native cool-season grass, Fescue (Festuca pratensis).

To stimulate flowering (Kucera and Ehrenreich 1962) and

encourage the spread of Indian grass into larger areas, we

experimented with a late-fall burn in 1997. Three 50 meter x

3 meter belt transects were established to facilitate counting

fertile stems before and after the treatment.

Results and Discussion

Daisy-leaved moonwort

The population responded favorably to both treatments,

prescribed fire and raking (23 plants in 1997 to 114 plants in

1999) (Figs. 1-4). The fire treatment showed a 35 percent

increase in plants and the raking treatment showed a 33

percent increase. The control has shown no increase.

0

50

100

1991 1994 1997

total # plants

# plants

trendline

0

20

40

60

80

1997 1998 1999

total # plants

# plants

trendline

burn

0

20

40

60

80

1991 1994 1996 1997

total # plants

# plants

trendline

0

20

40

60

80

1997 1998 1999

total # plants

# plants

trendline

Figure 1.—Number of Daisy-leaved moonworts

(Botrychium matricariifolium) before the 1998 spring fire.

Figure 2.—Number of Daisy-leaved moonworts

(Botrychium matricariifolium) after the 1998 spring fire.

Figure 3.—Number of Daisy-leaved moonworts

(Botrychium matricariifolium) before 1998 spring raking.

Figure 4.—Number of Daisy-leaved moonworts

(Botrychium matricariifolium) after 1998 spring raking.
118 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Figure 8.—Average number of fertile stems per 1-meter square in Indian

grass population (Sorgastrum nutans) before fire (1997) and after fire (1998).

0

500

1000

1500

2000

1991 1997 1998 1999

Total # rosettes

fertile

sterile

0

200

400

600

800

1000

1200

19891990199119921993199419951996199719981999

total # stems

# stems

infected

0

200

400

600

800

1000

1200

19891990199119921993199419951996199719981999

total # stems

# stems

infected

clip/rake

0

20

40

60

80

100

120

123

Belt Transect (50 m X 3m)

# fertile stems per 1-meter square

1997: Pre-burn

1998: Post-burn

Figure 5.—Number of fertile and stertile Bittercress

(Cardamine flagellifera) rosettes 1991-1999 in burned

portions of the population.

Figure 6.—Number of Goldenseal (Hydrastis canadensis) stems in subpopulation 1

from 1989-1999 with number of infected stems.

Figure 7.—Number of Goldenseal (Hydrastis canadensis) stems in subpopulation 2

from 1989-1999 with number of infected stems.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 119

Bittercress

A 27 percent increase in rosettes was observed from 1998 to

1999, all within the burned portion of the study site. In

addition, fertile rosettes showed a twofold increase (118

percent) after prescribed fire (Fig. 5).

Goldenseal

After our first treatment (clipping, etc.), a large reduction in

diseased stems was documented in 1992 (90 percent).

Subsequently, diseased stems in subpopulation 1 more than

doubled (110 percent increase) from 1993 to 1994 (two

years after our first treatment) (Fig. 6). However, the

incidence of disease again decreased in 1995 without

intervention. An additional 93 percent decrease in infected

stems in this subpopulation was observed from 1998 to 1999

following our second treatment (prescribed fire) in March

1998 (Fig. 7). Data indicates both treatments (clip and rake,

or burn) are effective management tools.

Indian Grass

Indian grass responded favorably to prescribed fire. Average

number of fertile stems per 1-meter square increased in all

three belt transects after treatment (Fig. 8). Transect 1 had

the highest increase in average number of fertile stems per

1-meter square (8.94 in 1997 to 99.73 in 1998). Transects 2

and 3 had an average of 6.92 and 4.08 fertile stems in 1997,

respectively. In 1998, the average fertile stems per 1-meter

square increased to 72.05 and 20.9 in transects 2 and 3,

respectively.

Conclusions

Although fires were historically uncommon in southern

Appalachian cove forests (Harmon 1982), prescribed fire has

been an effective management tool for some rare plant

populations in GSMNP. Dormant-season burning to reduce

leaf litter buildup was successful. Daisy-leaved moonwort

and Bittercress populations increased after fire was

experimentally introduced. Prescribed fire also reduced the

incidence of a fungal infection in Goldenseal.

Raking was an effective tool at the Daisy-leaved moonwort

and Goldenseal populations as well. Managing selected rare

plant populations with prescribed fire and/or raking will

continue to be a part of our Rare Plant Program.

Literature Cited

Anderson, K.L.; Smith, E.F.; Owensby, C.E. 1970. Burning

Bluestem range. Journal of Range Management. 23: 81-

92

Dix, R.L.; Butler, J.E. 1954. The effects of fire on a dry,

thin-soil prairie in Wisconsin. Journal of Range

Management. 7: 265-268.

Hadley, E.B.; Kieckhefer, B.J. 1963. Productivity of two

prairie grasses in relation to fire frequency. Ecology.

44: 389-395.

Harmon, M.E. 1982. Fire history of the westernmost

portion of Great Smoky Mountains National Park.

Bulletin of the Torry Botanical Club. 109: 74-79

Kucera, C.L.; Ehrenreich, J.H. 1962. Some aspects of

annual burning on central Missouri prairie. American

Midland Naturalist. 72: 142-147.

Robocker, C.W.; Miller, B.J. 1955. Effects of clipping,

burning, and competition on establishment and

survival of some native grasses in Wisconsin. Journal

of Range Management. 8: 117-121.

USFWS/OSA. 1997. Proposal to include Hydrastis

canadensis in Appendix II of the Convention on

International Trade in Endangered Species of Wild

Fauna and Flora (CITES). 10th meeting of the

Conference of the Parties, Harare, Zimbabwe.
120 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Over time fire has played many roles in the modification of

the earth’s landscapes. In the beginning there was only

“natural” fire, but it came from several sources (i.e.,

volcanoes, spontaneous combustion, lightning, etc.). Then

came a new source of fire that has drastically altered the

Earth’s landscapes - cultural fire. Some 100,000 years ago,

as modern man evolved from earlier homo species, there

was a seminal event in mans “rise to civilization” - it was his

controlled use of fire. It is now recognized that this

accomplishment was a primary factor that enabled

Neanderthal man to move from Africa into the colder reaches

of central Europe. Much later (e 12,000 years ago) this same

tool enabled modern man to cross Berengia (the land bridge

between Siberia and Alaska that was exposed as continental

glaciation lowered ocean levels by capturing much of the

Earth’s water) and populate the Americas. The original

people of the Americas were modern human beings whose

arrival brought the first taste of “un-naturalness” to our

landscape.

There is strong public sentiment in the East today to

establish (restore??) “natural” forest conditions. Accepting

the environmentalists stance that “natural” means without a

human (cultural) influence, restoring this condition is an

impossibility as it has never before existed with the current

complement of trees. The deciduous forest moved northward

into the Eastern landscape between five and eight thousand

years ago as the Wisconsin Ice event ended. However,

“man” had arrived several millennia earlier when boreal

forests (spruce, larch, jack pine) extended as far south as

Atlanta. His activities (especially setting fires) possibility

aided in the northward migration of trees. Thus our present

forest cover evolved in a culturally altered environment and

many trees exhibit traits that adapt them to cultural

modifications of the environment (e.g., cones that do not

open until burned).

If it could be argued that prior to 1492 Native Americans lived

“in harmony” with their environment and no effect on it, then

the existence of a “natural” environment at that time could be

valid. However, the intent of this presentation is to show that

this was not the case; that throughout the Holocene (last

10,000 years) mans’ activities have resulted in marked

modifications in the Eastern landscape to the extent that,

then or now, few places can claim to display “natural”

conditions.

There were four preColumbian cultures that occupied the

Eastern landscape prior to 1492. The original settlers were

Paleo-Indian (10,000-8,000 BC), hunter-gathers who used

fire to drive and trap the megafauna (wooly mammoth,

mastodons, etc.) that served as a primary food source.

These fires also maintained the open, grassy conditions

essential for these animals (closed forests do not support

grass). The following period -the Archaic (8,000-1,000 BC)

saw the beginnings of agriculture with fire essential to control

the aggressive native vegetation that threatened to close

over their fields. Recent research indicates that

landscape-scale fires of cultural origin were occurring in the

Southern Appalachians during this Period. Next came the

Woodland period (1,000 BC-800 AD) in which agriculture

was refined and fire was widely used to open the landscape

for hunting, travel and for protection. The Mississippian

Period (800-cl, 600 AD) brought PreColumbian time to a

close with Native American fire practices passed on to early

white settlers.

Ecologists now know enough about forest dynamics and tree

biology to model a close approximation of the “natural” forest

condition for Eastern forests. For our present complement of

trees, the “natural” forest condition would be a closed,

multi-canopied, forest composed of shade tolerant

hardwoods. The only conifer in the mix would be hemlock on

moist, cool sites. The highly productive “pioneer” species

characteristic of earlier stages of forest succession (the

change in species composition over time, from

light-demanding “pioneer” trees to shade tolerant “climax”

species) would occur only as widely scattered individuals or

in the rare areas where natural disturbances opened the

forest. The characteristic trees of today’s landscape (yellow

pines, yellow-poplar, oaks, sweetgum, ash, hickory, etc.) that

represent early- to mid-successional stages, would be

largely absent. Examples of the final stage of forest

succession (climax forest) may actually exist on a few sites

that have not been disturbed for several centuries thus

allowing succession to run its full course.

Shifts toward this condition are well underway on public

lands in the East. Sites that once supported pure yellow pine

or pine-hardwood stands are now converting to hardwoods,

and oaks are being replaced by red maple and black gum.

These shifts are due largely to 70+ years of highly effective

fire protection. The recent reductions in harvesting and

additions to wilderness (legislating protection from

disturbance in perpetuity) will further erode the productivity of

Eastern forests for both wildlife and forest products.

Accepting that this trend will continue, the widespread use of

prescribed fire offers the only hope for establishing

(maintaining) the early successional stages that contribute

significantly to the health and productivity of Eastern forests.

In Summary: Fire in the Evolution of the Eastern Landscape - A Timeline

Ed Buckner 1

1University of Tennessee (retired), Knoxville, TN 37919, ph: 423-

524-8329, email: ebuckner@utk.edu
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 121

Poster Abstracts
122 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Oak Seedling Response to Fire and Herbivory

A.S. Adams and L.K. Rieske

University of Kentucky, Department of Entomology, Lexington, KY 40546

We manipulated arthropod and mammalian herbivory levels on white oak seedlings using a combination of

insecticide applications and fencing, and employed a split-split-plot design to assess the impact of singleand multiple-year burns on seedling growth over a two year period. Herbivory levels increased over time on

all sites, but there was no significant difference in herbivore pressure on seedlings in single-year,

multiple-year, and non-burned plots. Insecticide treated seedlings (+I) suffered less herbivore pressure than

did non-insecticide treated seedlings (I)(P<0.0001), and mammalian herbivory was significantly reduced by

the presence of a fence (+F)(P<0.01). Seedling relative height growth, shoot elongation, diameter growth,

and specific leaf mass were greatest on once-burned sites, intermediate on twice-burned sites, and least on

non-burned sites. Bud expansion was the only seedling performance parameter unaffected by burn

treatment. We found differences in herbivory levels due to insecticide application, suggesting that the

arthropod component was the more significant aspect of overall herbivory levels, but this herbivory did not

impact seedling performance, and seedling performance parameters were more closely linked to

mammalian herbivory. These results suggest that prescribed fire may be a practical component of a

management program designed to enhance oak regeneration.

Prescribed Burning: Effects on Forest Bird Populations

Vanessa L. Artman

Department of Evolution, Ecology, and Organismal Biology, 1735 Neil Avenue,

Ohio State University, Columbus, OH 43210

Prescribed burning is being applied on an experimental basis in southern Ohio to restore and maintain

oak-dominated forests. Effects of prescribed burning on bird communities have been given little

consideration in previous research, but are of concern because long-term declines have been observed for

many bird species within the region. My research addresses short-term changes in population levels and

nesting productivity of forest bird species. Bird species diversity did not change as a result of burning, but

population levels of three species, the hooded warbler, ovenbird, and worm-eating warbler, declined in

burned areas. As expected from the reduced population levels, fewer active nests were found in burned

areas than in unburned areas, but there were no differences in rates of nesting success between treatments.

Predation of nest contents was the primary factor influencing the likelihood of nesting success. A wide

variety of nest predators inhabit the forests, including snakes, squirrels, raccoons, and other birds, and their

foraging tactics are so variable that nest predation events tend to be unpredictable. Prescribed burning did

not appear to affect the array of nest predators in the community or the susceptibility or exposure of eggs or

nestlings to these predators.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 123

Ecological Effects of Fire on Xerophyllum asphodeloides,

a Rare Appalachian Lily

Norman A. Bourg and Douglas E. Gill

Department of Biology, University of Maryland, College Park, MD 20742

Fire’s role as a biotic disturbance agent in Appalachian forests has been historically understudied. We

performed a fire/canopy removal field experiment to determine the impact of these factors on survival and

reproduction of turkeybeard (Xerophyllum asphodeloides), a rare lily associated with Appalachian mixed

pine-oak forests. Turkeybeard is endangered or rare in portions of its range and is in the U.S. Center for

Plant Conservation’s National Endangered Plant Collection. It is also thought to be fire-dependent; yet, there

has been no experimentally validated ecological linkage to fire. An immense flowering response to our

treatments occurred in the second and third post-manipulation years. Mixed-model ANCOVA analyses

yielded a significant positive effect of fire on both the numbers of fertilized flowers and seeds. A significant

positive effect of canopy removal on these variables was also found. Total number of flowers per

inflorescence did not differ between control and treated plants. Inflorescences from another nearby

undisturbed, shaded population showed virtually no fertilization or seed production. Many plants exhibited a

cost of reproduction by partial vegetative dieback, but little complete mortality occurred. This suggests that

fire is important for turkeybeard conservation, and also has implications for the study of disturbance regimes

in Appalachian forests.

A Shelterwood-burn Technique for Regenerating Productive

Upland Oak Sites

Patrick Brose1 and David Van Lear 2

1

Research Forester, USDA Forest Service, Northeastern Research Station, Forestry Sciences Laboratory,

Irvine, PA 16329, 2

Bowen Professor of Forestry, Clemson University, Department of Forest Resources,

Clemson, SC 29634

Regenerating oak stands on productive upland sites is widely recognized by foresters as a major problem in

hardwood management. Recent research indicates that oak regeneration is a more resilient sprouter

following surface fires than its primary competitors on these sites if burning occurs 3 to 5 years after a partial

overstory harvest. This combination of cutting followed by fire (the shelterwood - burn technique) mimics

natural disturbances that have occurred in eastern North America for millennia and appears to be a viable

approach to regenerating oaks on productive upland sites. This poster presents silvicultural guidelines for

applying the shelterwood - burn technique on appropriate sites and discusses its benefits for private

landowners and resource professionals.
124 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Effects of Fire, Deer Herbivory, Seed Predation, and Canopy Gaps on Forest

Regeneration: A Large-scale Experimental Approach

Rachel J. Collins and Walter P. Carson

Department of Biological Sciences, University of Pittsburgh, Pittsburgh, PA

Oaks have been a dominant group and readily regenerating in eastern deciduous forests for thousands of

years. Interestingly, since the 1960s, oaks have failed to regenerate in many areas. This pattern is one of a

number of symptoms characteristic of recent changes in forest understories (i.e. the decline in diversity of

wildflowers and tree seedlings; and the increase in maple abundance). We are currently investigating the

underlying mechanisms that control tree species regeneration in eastern forests using a large-scale

experimental approach to test four prevalent hypotheses. We have experimentally manipulated fire, deer,

seed predation and canopy gaps in 128, 400m2

 research plots in West Virginia and are monitoring the

survival, growth and recruitment of the 28,000 seedling, sapling and canopy trees within these research

plots. The fires were intense and topkilled up to 100% of small saplings and up to 60% of large saplings;

however canopy tree mortality was only 4% and not different from controls. Fires strongly influenced the

species composition of emergent seedlings. Fire by deer, and gap by deer interactions are emerging as

important factors affecting growth and survival of seedlings. This study is just beginning to elucidate the

underlying mechanisms that drive forest regeneration.

Effects of Shelterwood Harvesting and Prescribed Fire

on Regeneration Structure

Wade Dorsey1 and Mary Ann Fajvan2

1

Doctoral Candidate, 2Associate Professor, West Virginia University, Division of Forestry, Morgantown,

WV 26506

In 1993, four, 10 acre Appalachian hardwood stands received a shelterwood seed cut. Preharvest overstory

species consisted primarily of yellow poplar (40-50% of basal area) and oaks (30% of basal area);

understory composition was dominated by red maple. Post-harvest basal areas averaged 70ft2

/a with oaks

comprising the majority of the seed trees. After 6 growing seasons, black cherry, red maple and sassafras

seedlings dominated the regeneration. Because regeneration composition did not reflect the pre-harvest

overstory, a prescribed fire was conducted in November 1999 primarily to reduce the density of black cherry

and red maple and to promote oak species. The effects of fire on herbaceous vegetation and soil nutrients

are also being examined.

In the summer of 1999, the 10-acre treatment blocks were divided into two, 5-acre treatments: burned and

unburned. In each 5-acre block, nine, milacre sampling plots were established and all woody and

herbaceous vegetation <1" dbh was counted and height recorded. An additional milacre was established to

measure only tree seedlings. All plots will be remeasured in the spring and summer of 2000 and 2001.

Soil samples were collected at 3 locations, adjacent to six of the vegetation sample plots, immediately prior

to and after burning. Soil samples will be collected two additional times in spring and summer 2000. Litter

and mineral soil samples will be analyzed for total N. P. K, Mg, Ca, and S. Mineral soil samples will also be

analyzed for exchangeable or available N. P. K, Mg, Ca, and S. CEC, % BS, bulk density and pH.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 125

Prescribed Fire Effects on Foliar Nutrients, Photosynthesis, and Growth

of Ridgetop Maple and Oak Seedlings

N.L. Gilbert, M.A. Arthur, S.L. Johnson, S.K. Gleeson and B.A. Blankenship

University of Kentucky, Lexington, KY

Forest management with low-intensity prescribed burns aims to promote oak regeneration in upland forests

of the Cumberland Plateau, where sixty years of fire suppression coincide with increasing red maple

abundance. Our objective was to determine whether a single fire influences the ecophysiology of red maple,

chestnut oak, and scarlet oak seedlings during the first year postburn. On a burned (March 1997) and an

unburned ridgetop site, we measured foliar nutrients (N. P. K, Mg, and Ca) monthly during the 1997 growing

season, photosynthesis at five light intensities (0, 100, 400, 1000, 1500 uEm-2s-1) in July, 1997, and stem

diameter and height in May, 1997, and March and May, 1998. For all species, foliar N. P and K, net

photosynthetic rates, and diameter and height relative growth rates were significantly higher on the burned

site than the unburned site. Fire had little effect on the relative ecophysiology among species on the burned

site compared to the unburned site. These data suggest that single low-intensity prescribed burns will not

benefit the regeneration of oaks by enhancing their ecophysiological performance over that of red maple.

Chilton Creek: A Case Study of Landscape Fire Effects

George W. Hartman1 and Blane Heumann2

1

Fire Ecologist, Missouri Department of Conservation, 1110 S. College Ave., Columbia, MO 65201, 2Lower

Ozarks Preserves Manager, The Nature Conservancy, Missouri Ozarks Office, P.O. Box 960, Van Buren,

MO 63965

The Chilton Creek Management Area is 5,657 acres located in Carter County, Missouri on the Ozark

Plateau. Owned and managed by The Nature Conservancy, a 2,500-acre portion is divided into 5 burn units

with a fire return interval for the various units from annual to 4 years The major objective of the studies is to

document the effects of landscape-scale prescribed burning on the naturally occurring communities of the

Ozark woodlands. Fuel loading, fire behavior, fire intensity and coverage measurements are also being

recorded to relate fire activity to resource response. Prescribed burning was initiated spring, 1998.

The study to document fire effects on the herbaceous and woody components is being conducted

cooperatively with the Missouri Department of Conservation. Nested plots (250) and sampling protocols

mimic an adjacent long term study of timber harvesting techniques called the Missouri Ozark Forest

Ecosystem Project (MOFEP). Prescribed fire management will be compared to even aged timber

management, uneven aged management and controls (no active management)

Other studies being conducted with the restoration burning include stream hydrology and biota, differential

mortality of young woody species, breeding bird populations, herpefauna, lichens, invertebrates including

land snails and Orthoptera, soils, geology, and ecological land classifications.
126 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Ethnoecology of Fire: An Experimental Approach in the Ohio Valley

Erin B. Hines1, Jan Salick1, Elaine Kennedy Sutherland2, Todd F. Hutchinson3 and Steve Sutherland4

1

Department of Environmental and Plant Biology, Ohio University, Athens, OH 45701, 2USDA Forest Service,

Rocky Mountain Research Station, Forestry Sciences Laboratory, PO Box 8089, Missoula, MT 59807,

3

USDA Forest Service, Northeasterm Research Station, 359 Main Road, Delaware, OH 43015, 4USDA

Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, P.O. Box 8089, Missoula, MT

59807

Native Americans used fire to manipulate nature and directly benefit their survival. Certain plant species,

many of which were useful to Native Americans as sources of food, fiber, dye, medicine, and game browse,

are adapted to survive and even thrive in post-burn environments. Evidence suggests that Native Americans

intentionally set fires to encourage growth and survival of such useful species. Data from a 5-year study

conducted by the U.S. Forest Service in Southeastern Ohio to track the long-term effects of prescribed fire

on forest composition were used to track changes in useful plant abundance under certain fire regimes.

Preliminary analyses of pre-burn data establish covariates between burn treatments and moisture indices.

Split-plot block ANOVAs of the 1999 post-burn data suggest an increase in many fern and graminoid

species, along with an increase in Vaccinium and early-successional tree species. The results from this

study have implications for modern forest management and global conservation. Fire increases diversity in

both botanical and zoological systems by initiating patches of early succession in plant communities, thus

increasing resources available for ecosystem interactions and possible human use. By encouraging the

growth of both timber and non-timber forest products, resource managers can move towards more

multiple-use forest management.

Managing Red-Cockaded Woodpeckers (Picoides borealis) Affects

Breeding Bird Communities of Pine-Oak Forests in Southeastern Kentucky

Martina Hines

Kentucky State Nature Preserves commission

Breeding bird communities were surveyed on the Daniel Boone National Forest, Kentucky, using point

counts, in mid May, late May and early June 1994 and 1995. Twenty-eight stands of Appalachian pine-oak

forest were surveyed, of which 14 were managed for red-cockaded woodpeckers (RCWs). Management

involved removal of midstories (2.5 to 17.5 cm dbh) and repeated prescribed burns. Results indicate that

these management practices appear to have effects similar to that of timber harvests on species

composition and abundance of breeding birds. Prescriptions for RCWs were beneficial to several species

associated with early successional and edge habitat, including at least some neotropical migrants whose

numbers are decreasing in eastern North America (e.g., indigo bunting, hooded warbler, and prairie warbler).

Management for RCWs appears to decrease habitat quality for other neotropical migrants, particularly those

associated with forest interiors.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 127

Disturbance History of an Old-Growth Forest in Southeastern Ohio

Brian C. McCarthy and Darrin L. Rubino

Department of Environmental and Plant Biology, Ohio University, Athens, OH 45701

We performed a dendroecological investigation of Dysart Woods, an old-growth forest in southeastern Ohio,

to determine stand age and the historic disturbance regime. The 23 ha woods is dominated by white oak,

beech, and sugar maple. Because Dysart Woods is a preserve, our sample size was limited to ten white oak

samples that yielded a 374-yr chronology. Of the samples with pith, it appears that the trees originated under

closed canopy conditions. From 1625-1850, ring widths were consistently small and below the overall mean

of 1.5 mm; this was followed by a nearly 100-yr period of above-average growth before leveling off in the last

50 years. Using a 10-yr running median technique, we detected 2.42 release events per tree per 100 years.

This is consistent with other old-growth studies in the region and characteristic of a gap disturbance regime.

With few exceptions, Resettlement fire is not recorded in a full basal slab from Dysart Woods. However,

burning during the 1800s (period of settlement) was prolific and occurred every few years before diminishing

in the early 1900s. While fire may have been an important ecological factor in these woods, it does not

appear to have influenced the origin of white oak.

Response of Pin Cherry (Prunus Pensylvanica L.F.) to Fire,

Canopy Disturbance, and Deer Herbivory on the Westvaco Wildlife

and Ecosystem Research Forest

David W. McGill1, Rachel J. Collins 2, and Walter P. Carson2

1

Research Scientist, Westvaco Corporation, Appalachian Forest Research Center, Rupert, WV 25984, 2

Doctoral Candidate and Associate Professor, respectively, Department of Biological Sciences, University of

Pittsburgh, A234 Langley Hall, Pittsburgh, PA 15260

A study was established to assess the impact of fire, canopy disturbance, and deer herbivory on the

germination and development of pin cherry (Prunus pensylvanica L.f.) in Appalachian hardwood stands

located on the Westvaco Wildlife and Ecosystem Research Forest in Randolf County, West Virginia. In each

of four stands, two blocks with four, 400 square meter treatment plots were established. One of four

treatments were randomly applied to each treatment plot: 1) fence, gap; 2) fence, no gap; 3) no fence, gap;

and 4) no fence, no gap. In each treatment plot, five one-square meter regeneration plots were randomly

placed and permanently monumented. All four stands were burned in spring 1999. Over the course of the

1999 growing season, four separate regeneration measurements were made at approximately one-month

intervals. Pin cherry germination showed a large amount of variation from stand to stand. Average initial pin

cherry densities ranged from 23,500 to 63,000 seedlings per hectare (SPH) following prescribed fire. By the

end of the growing season, seedling densities ranged from a low of 5,250 SPH in the No Fence/Gap

treatment to 43,250 SPH in the Fence/Gap treatment. These differences were principally a function of deer

herbivory. Average height in the Fence/Gap treatment was 12.6 cm greater than the next tallest treatment

average—the Fence/No Gap treatment. The tallest pin cherries in the fenced treatments exceeded one

meter in height.
128 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274

Prescribed Fire and Herbicides Modify Soil Processes During Barrens

Ecosystem Restoration

C.C. Rhoades, T. G. Barnes, and B.E. Washburn

University of Kentucky, Department of Forestry, Lexington, KY 40546

Prescribed fire has become a common tool of natural area managers for removal of invasive species and

maintenance of barrens grassland communities. Certain target species however, resist fire and may require

additional removal treatments. An effective option for removal of Festuca arundinacea (tall fescue) from

areas that previously supported native warm season grasses combines spring prescribed fire and

post-emergent herbicides. We studied changes in soil N and C dynamics following prescribed fire and

herbicide application in a remnant barrens in westcentral Kentucky. Fire and herbicide effects were

compared in a 2 way factorial design with 5 replicates. Fescue cover decreased and bare soil increased

from near 88 and 2% in no-burn/noherbicide control plots to 12 and 28% in burned/herbicide plots. During

the month following the fires, soil N movement measured with in situ exchange resins was 1.4-fold higher in

burned compared to unburned soils. Net N mineralization was 2.8-fold higher in burned compared to

unburned soils. Burned/herbicide soils mineralized more N than non-herbicide soils. Fire increased soil

temperature and decreased soil moisture and CO2 efflux. Removal of non-native plants modify chemical,

physical and biologic soil conditions that control availability of plant nutrients and influence plant species

performance and community composition

Experimental Studies of Oak Seed Responses to Fire Temperature

in Appalachian Mixed-oak Litter

Cynthia L. Riccardi and Brian C. McCarthy

Department of Environmental and Plant Biology, Ohio University, Athens, OH 45701

The relationship between wildland fire intensity, fire temperature, and the response of plant and animal

populations is complex. Few studies have examined the relationship between fire temperature at fine spatial

scales and oak seed germination. To examine temperature patterns, we exposed aluminum tags painted

with temperature-sensitive paints (38-427°C) and placed them at –1, +2, and +12 cm relative to the soil

surface, in 1 x 1 m grids with 20 cm spacing distances, and burned in oak litter. Belowground (<30°C) and

12 cm (range: 150-225°C) sensors were spatially homogeneous across the experimental burn plots.

Sensors in the leaf litter experienced the hottest and most variable temperatures (95-375°C) and greatest

spatial variability. A subsequent study of 432 Quercus prinus seeds varied seed position (above- or

below-ground), temperature (control, 38, 66, 150, 204, and 260°C), and exposure time (5 or 15 min). Time

and temperature were highly significant (P<0.001). Seed exposure to temperatures above 150°C for more

than 5 min yielded decreased germination. Temperatures at 204°C for 15 min usually resulted in complete

mortality. Caching of Q. prinus seeds by gray squirrels may potentially increase seed survival during wildland

fires. Further studies are underway to evaluate temperature and its effect on weevil (Curculio) populations.
Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 129

Ground Beetle Responses to Prescribed Burning in Southern Ohio’s

Hardwood Forests

Chris Stanton, Foster F. Purrington, and David J. Horn

The Ohio State University, Department of Entomology, Columbus, OH 43210

The Forest Service has administered prescribed fire to hardwood forests in southern Ohio since 1996 in

order to study the ecological response of the mixed-oak community to prescribed burning. One component

of the project involved studying the effects of the fires on forest floor invertebrates. The primary objective of

this study was to monitor ground beetle populations for any significant changes that may indicate critical

effects of fire on the forest floor community.

Four replicates of mature, mixed-oak forests were subjected to annual spring burns, occasional spring

burns, and no burns (control). Ground beetles were collected via pitfall traps established within each

treatment block and operated from May to October each year.

Results from five years of monitoring do not indicate any significant effects of prescribed fire on ground

beetle abundance, species richness, or diversity. Because the fires did not significantly affect ground beetle

populations, prescribed burning should be considered a safe management tool for restoring and maintaining

oak forests without devastating the forest floor community.

Monitoring Scarab Populations Exposed to Prescribed Fire in Mixed-oak

Forests of Southeast Ohio

A.E. Smith and D.J. Horn

Department of Entomology, The Ohio State University, Columbus, OH 43210

Scarabaeidae was selected to determine the effects of surface fire on the distribution and abundance of

insect populations. Subfamilies Geotrupinae and Scarabaeinae were expected to show impact of fire,

because they exploit resources, such as leaf litter and fungi, that are consumed by fire. Twelve 60-ha. study

blocks were selected in 80- 120 year old mixed-oak forest according to similar exposure, soil type, elevation,

and vegetation composition. Six blocks each were located in Vinton and Lawrence County, Ohio. Each block

was equipped with a transect of twelve, 11-cm dia. unbaited pitfall traps. A pre-burn baseline survey was

completed in 1995, and two blocks per site were burned in April 1996; one block was burned annually

1997-1999, and the other was burned in 1999. Three species were selected to model burning effects:

Ateuchus histeroides, Onthophagus striatulus, and Geotrupes.splendidus. A multivariate analysis of

repeated measures revealed no measurable impact of fire; however, significant yearly variation did exist.

Data show a high interplot variation, which suggests that species could be habitat specific. This study yields

insight into the diversity and distribution of Scarabaeidae, and whether forest plots can be adequately

replicated.
Printed on Recycled Paper

Yaussy, Daniel A., comp. 2000. Proceedings: workshop on fire, people, and the

central hardwoods landscape; 2000 March 12-14; Richmond, KY. Gen. Tech.

Rep. NE-274. Newtown Square, PA: U.S. Department of Agriculture, Forest

Service, Northeastern Research Station. 129 p.

Contains 18 papers and 16 poster abstracts on the history of fire, fire ecology, fire

and ecosystem management, and fire and the future presented at the workshop on

fire, people, and the central hardwoods landscape.

Keywords: native burning, prescribed fire, prescribed burning, oak, mixed-oak, oakhickory, barrens, ridgetop-pine, soil microbes, rare plants
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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