United States

Department of 

Agriculture

Forest Service

Northern 

Research Station

General Technical

Report NRS-P-1

Fire in Eastern Oak Forests:

Delivering Science to Land Managers

Proceedings of a Conference

November 15-17, 2005

Fawcett Center 

The Ohio State University, 

Columbus, Ohio
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Acknowledgments

The Steering Committee thanks Sarah Sieling, Robin Dever, and Laura Seeger, Department 

of Conference Management and Professional Development, The Ohio State University, 

for logistical support. We also thank the many reviewers for improving the manuscripts. 

Generally, there were two anonymous reviewers for each manuscript, one from the land 

management and one from the scientific community. Anantha Prasad created and maintained 

the conference website. Mary Boda provided administrative support. Financial support was 

provided by the Northeastern Area, State and Private Forestry, Southern Region, and Joint 

Fire Science Program, The Nature Conservancy’s Global Fire Initiative, and Lion Apparel. 

In-kind support was provided by The Ohio State University, School of Natural Resources, 

Northeastern Research Station, and Ohio Department of Natural Resources.

The findings and conclusions of each article in this publication are those of the individual 

author(s) and do not necessarily represent the views of the U.S. Department of Agriculture or 

the Forest Service. All articles were received in digital format and were edited for uniform type 

and style. Each author is responsible for the accuracy and content of his or her paper.

Abstract

Contains 20 papers and 36 poster abstracts presented at a conference on fire in oak forests 

of the Eastern United States that was held at the Ohio State University, Columbus, Ohio, on 

November 15-17, 2005.
Fire in Eastern Oak Forests:

Delivering Science to Land Managers

Proceedings of a Conference

November 15-17, 2005

Fawcett Center, The Ohio State University, Columbus, Ohio

Edited by

Matthew B. Dickinson

USDA Forest Service, Northern Research Station

Forestry Sciences Laboratory, Delaware, Ohio 43015

Steering Committee

Mike Bowden, Ohio Department of Natural Resources

Pat Brose, USDA Forest Service, Northern Research Station

 Matthew Dickinson (Chair), USDA Forest Service, Northern Research Station

Rex Mann, USDA Forest Service, Daniel Boone National Forest

Roger Williams, The Ohio State University, School of Natural Resources

Alan Zentz, USDA Forest Service, Northeastern Area, State and Private Forestry

Sponsors

USDA Forest Service, Northeastern Area, State and Private Forestry

USDA Forest Service, Southern Region

Joint Fire Sciences Program

The Ohio State University

USDA Forest Service, Northern Research Station

Ohio Department of Natural Resources

The Nature Conservancy’s Global Fire Initiative

Lion Apparel
	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Contents

Historical Perspectives on Eastern Oak Forests

The Paleoecology of Fire and Oaks in Eastern Forests ..............................................................2

William A. Patterson III

Fire Scars Reveal Variability and Dynamics of Eastern Fire Regimes ....................................20

Richard P. Guyette, Daniel C. Dey, Michael C. Stambaugh, and Rose-Marie Muzika

Understanding the Evidence for Historical Fire Across Eastern Forests................................40

Charles M. Ruffner

Oak Composition and Structure in the Eastern United States..................................................49

W. Keith Moser, Mark Hansen, Will McWilliams, and Ray Sheffield

Why Sustain Oak Forests?...........................................................................................................62

David Wm. Smith

Vegetative and Forest Floor Responses to Fire

Ecological and Ecophysiological Attributes and Responses to Fire in Eastern 

Oak Forests....................................................................................................................................74

Marc D. Abrams

Research Efforts on Fuels, Fuel Models, and Fire Behavior in Eastern 

Hardwood Forests.........................................................................................................................90

Thomas A. Waldrop, Lucy Brudnak, Ross J. Phillips, and Patrick H. Brose

Soil, Fire, Water, and Wind: How the Elements Conspire in the Forest Context...................104

Ralph E.J. Boerner

Responses of Oak and Other Hardwood Regeneration to Prescribed Fire: 

What We Know as of 2005 ..........................................................................................................123

Patrick H. Brose, Thomas M. Schuler, and Jeffrey S. Ward

Fire and the Herbaceous Layer of Eastern Oak Forests..........................................................136

Todd Hutchinson

Fire—Fauna Interactions

Do Fire and Insects Interact in Eastern Forests?.....................................................................152

Lynne K. Rieske-Kinney

Does Fire Affect Amphibians and Reptiles in Eastern U.S. Oak Forests?.............................158

Rochelle B. Renken

Conservation Implications for Neotropical Migratory and Game Birds in Oak-hardwood 

Stands Managed with Shelterwood Harvests and Prescribed Fire ........................................167

J. Drew Lanham, Patrick H. Brose, and Patrick D. Keyser

Influence of Fire on Mammals in Eastern Oak Forests............................................................180

Patrick D. Keyser and W. Mark Ford
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Obstacles and Solutions to Using Prescribed Fire in Oak Forests

 Prescribed Fire: What Influences Public Approval?...............................................................192

Sarah M. McCaffrey

Top Ten Smoke Management Questions for Fire in Eastern Oak Forests .............................199

Joseph J. Charney, Ann L. Acheson, and Andrea Stacy

Resistance of Eastern Hardwood Stems to Fire Injury and Damage .....................................210

Kevin T. Smith and Elaine Kennedy Sutherland

Fire and Invasive Exotic Plant Species in Eastern Oak Communities: 

an Assessment of Current Knowledge......................................................................................218

Cynthia D. Huebner

Rx Fire Laws: Tools to Protect Fire: the ‘Ecological Imperative’............................................233

Dale Wade, Steven Miller, Johnny Stowe, and James Brenner

Growing a Burning Program: Challenges and Opportunities.................................................263

Rex Mann

Poster Abstracts

Overstory and Seedling Response to Repeated Fires in Eastern Kentucky .........................268

Heather D. Alexander, Mary Arthur, David Loftis, and Stephanie Green

Thermocouple Probes in Surface Fires.....................................................................................269

Anthony S. Bova and Matthew B. Dickinson

Wildfire Hazard Mitigation Treatments Applied Above Ground Impact Soil 

Ecological Properties in Oak Forest Ecosystems....................................................................270

Jennifer A. Brinkman and R.E.J. Boerner

Fire Research in the Pine Barrens of New Jersey....................................................................271

Kenneth Clark, John Hom, Nick Skowronski, Yude Pan, Steve Van Tuyl, and Warren Heilman

Effects of Fuel Reduction Treatments on Soil Nitrogen Transformation in 

the Southeastern Piedmont........................................................................................................272

T. Adam Coates, Victor B. Shelburne, Thomas A. Waldrop, Bill R. Smith, and Hoke S. Hill, Jr.

Use of Wildland Fire in Great Smoky Mountains National Park..............................................273

Dana Cohen, Beth Buchanan, Bob Dellinger, Leon Konz, and Mark Taylor

Hardwood Stem Injury and Mortality in Surface Fires.............................................................274

Matthew B. Dickinson and Anthony S. Bova

Vegetation Response to Prescribed Fire Across a Moisture/Productivity Gradient 

in the Southern Appalachians....................................................................................................275

Katherine J. Elliott and James M. Vose

Oaks in Presettlement Fire Landscapes of the Southeast......................................................276

Cecil Frost

Effects of Prescribed Burning on Invasibility by Nonnative Plant Species in 

the Central Hardwoods Region..................................................................................................277

Lance Glasgow and Glenn Matlack
	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Fuel and Fire Dynamics in Eastern Mixed-Oak Forests...........................................................278

John B. Graham and Brian C. McCarthy

Using Prescribed Fire and Mechanical Thinning to Improve the Regenerative 

Potential of Oak Species.............................................................................................................279

Scott Holevoet and Charles Ruffner

Thinning and Burning Treatments Influence Within-Canopy Leaf Traits in Seven 

Hardwood Species in Ohio.........................................................................................................280

Jyh-Min Chiang and Kim J. Brown

Estimating Fire Behavior from Overhead Imagery in Eastern Oak Forests...........................281

Robert Kremens, Anthony Bova, Matthew Dickinson, Jason Faulring, and Shari McNamara

Effects of Silvicultural Treatments on Acorn Weevil Occurrence in Mixed-Oak Forests .....282

Jeffrey A. Lombardo and Brian C. McCarthy

Oak and Maple Stump Sprout Dynamics in Response to Thinning and Burning 

Treatments in Southern Ohio .....................................................................................................283

Robert P. Long, David K. Apsley, and Joanne Rebbeck

Impacts of Repeated Prescribed Fires on Fuels in a Central Appalachian 

Hardwood Forest.........................................................................................................................284

Jessi E. Lyons, Elizabeth Loucks, Mary A. Arthur, and David Loftis

Factors Influencing Wildfire Occurrence and Distribution in Eastern Kentucky..................285

John K. Maingi and Mary C. Henry

Hardwood Regeneration Following Prescribed Fire and Thinning in Mixed-Oak 

Forests of Southeastern Ohio....................................................................................................286

Brian C. McCarthy and Matthew A. Albrecht

Soil Respiration Responses to Temporal, Topographic, and Silvicultural Factors in 

a Southeastern Ohio Forest .......................................................................................................287

Dawn R. McCarthy and Kim J. Brown

Changes in the Stand Structure and Species Composition of Oak-dominated 

Communities Following a Prescribed Burn..............................................................................288

Virginia L. McDaniel, Michael A. Jenkins, and Robert N. Klein

Experimental Evaluation of Fire History Reconstruction Using Dendrochronology 

in White Oak (Quercus alba).......................................................................................................289

Ryan W. McEwan, Todd Hutchinson, Robert Ford, and Brian C. McCarthy

Impact of Fire on Soil Resource Patterns in Mixed-Conifer Forests in the Southern 

Cascade Range of Northern California......................................................................................290

Jessica Miesel, Carl Skinner, and Ralph Boerner

Avian Community Responses to Prescribed Burns and Shelterwood Harvests ..................291

Donald B. Miles, Henry Streby, and Dan Weber

Rehabilitation of an Upland Hardwood Forest Using Prescribed Fire and Herbicide ..........292

Rebecca Montgomery, Jeff Earl, Chris Stuhlinger, Matthew Pelkki, and Kyle Cunningham

Altered Disturbance Regimes: Demise of Fire in the Eastern United States.........................293

Gregory J. Nowacki and Robert A. Carr
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 

Lightning-initiated Growing Season Fires Characterized Oak-Dominated Ecosystems

 of Southern Ohio .......................................................................................................................294

Sheryl M. Petersen and Paul B. Drewa

Use of Temperature-Sensitive Paints as an Index of Heat Output From Fire........................295

Joanne Rebbeck, Daniel Yaussy, Louis Iverson, Todd Hutchinson, Robert Long, Anthony Bova, 

and Matthew Dickinson

Impact of Fire, Silvicultural Manipulation, and Microsite Conditions on Woody 

Debris Decay Dynamics in a Mixed-Oak Forest .......................................................................296

Darrin L. Rubino, Lindsey E. Hesch, and Brian C. McCarthy

Effects of Forest Management on Soil Seedbank Species Composition in a Mixed-Oak 

Forest in Southeastern Ohio ......................................................................................................297

Lisa R. Schelling and Brian C. McCarthy

Oak-Grassland Restoration Demonstration Area Project .......................................................298

Leslie M. Smith and James R. McCoy

Integrated Upland Hardwood Research in Arkansas’ Boston Mountains..............................299

Martin A. Spetich

Assigning Fire Regimes on the Monongahela National Forest, West Virginia......................300

Melissa Thomas-Van Gundy

Influence of Prescribed Fire on Stem Girdling and Mortality..................................................301

Jeffrey S. Ward and Patrick H. Brose

Putting Fire on the Ground with Prescribed Burning Associations.......................................302

John R. Weir, Terrence G. Bidwell, and Chris M. O’Meilia

Fire Research on The Raccoon Ecological Management Area...............................................303

Daniel A. Yaussy, Todd Hutchinson, Matthew B. Dickinson, and Joanne Rebbeck
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 

Historical Perspectives 

on Eastern Oak Forests
	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

THE PALEOECOLOGY OF FIRE AND OAKS IN EASTERN FORESTS

William A. Patterson III1

Abstract.—Oaks (Quercus spp.) currently dominate eastern deciduous forests, but are widely perceived 

as declining, with regeneration inadequate to perpetuate many stands. Most stands regenerated following 

fire in the 19th and early 20th centuries, and a lack of recent fire is viewed as contributing to the shortage 

of sapling and pole-size stands. But paleoecological studies provide conflicting evidence for the role of 

fire in the long-term maintenance of oak forests. Here I describe the methods used in reconstructing past 

vegetation and fire regimes, review the results of previous studies, and present new results for sites from 

Virginia to Maine. Oaks have dominated mixed mesophytic forests in western Virginia for more than 

6,000 years, with sedimentary charcoal levels suggesting a fire regime dominated by infrequent, light 

surface fires. The arrival of European settlers and a presumed increase in fire activity had little effect on 

oak abundance. At a higher elevation on more xeric soils, increased fire with settlement caused a shift 

from oak to pine dominance. On Long Island, NY, oaks have dominated xeric soils for thousands of 

years, but with more fire than in western Virginia. However, a dramatic increase in fire activity with 

settlement increased the importance of pines relative to oaks in southeastern Massachusetts’ outwash 

plains. On the Maine coast, where oaks have been minor component of the vegetation for thousands 

of years, fires appear to have caused slight increases in oak importance both before and especially since 

European settlement. These results suggest that, depending on the landscape context, fire can favor or select 

against oaks, and that managers should carefully consider how fires will interact with climate, topography, 

and other factors before prescribing fire as a solution to the current lack of oak regeneration. It is likely 

that burn severity and fire return intervals, as they impact both oaks and their potential competitors, 

will determine whether or not individual oak stands will benefit from the reintroduction of fire.

interface areas, browse and kill seedlings and saplings. 

Late 20th-century fire suppression is blamed for the lack 

of regeneration in many stands. Tirmenstein (1991) 

citing several studies, observed: 

Fire has played an important role in deciduous 

forests of the eastern United States. Evidence 

suggests that most oaks are favored by a regime 

of relatively frequent fire. Many present-day 

oak forests may have developed in response to 

recurrent fire. Declines of oak forests have been 

noted throughout much of the East and are often 

attributed to reduced fire frequency.

Detailed information on the effects of fire on many oak 

species is available through the Fire Effects Information 

System. Most are viewed as being favored, at least in the 

regeneration stage, by fire. Tirmenstein (1991) noted, in 

discussing the fire ecology of white oak: “it is unable to 

regenerate beneath the shade of parent trees and relies 

on periodic fires for its perpetuation. The exclusion of 

fire has inhibited white oak regeneration through much 

of its range.” Crow (1988) argued that frequent fires are 

required to maintain northern red oak (Quercus rubra), 

INTRODUCTION

With at least 36 species, one or more of which occur 

in every state east of the Great Plains (Samuelson and 

Hogan 2006), oaks (genus Quercus) are the preeminent 

trees of the eastern forest. Twenty-one species are at 

least locally important as timber species (Burns and 

Honkala 1990). All, and especially those of the white 

oak (Leucobalanus) group, provide mast for game and 

nongame species of wildlife. Species not important for 

timber (e.g., the scrub oaks as a group) provide essential 

food and cover for several species of Lepidoptera (moth) 

larvae that are rare in at least portions of their ranges 

(Wagner et al. 2003). Oaks dominate in a variety 

of habitats, from rich cove forests of the southern 

Appalachians to dry ridgetops and sand plains of New 

England and the Atlantic Coastal Plain.

Throughout the range of oaks there is a widely held 

belief that regeneration is inadequate to perpetuate 

existing stands. Abundant deer, especially in urban 

1

Forestry Program, University of Massachusetts, 214 

Holdsworth Natural Resources Center, Box 34210, Amherst, 

MA 01003, 413-545-1970, email: wap@forwild.umass.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

and that increased fire frequency has caused, in some 

cases, the replacement of mesic hardwoods (e.g. sugar 

maple, Acer saccharum) by red oak. Conversely, reduced 

fire frequency is resulting in the replacement of northern 

red oak by mesic hardwoods in many areas of the Lake 

States and Northeast (Crow 1988).

The important role of fire in the development of oak 

stands during the historic period, i.e., roughly since 

the arrival of European settlers on the North American 

continent, is well documented (see Abrams et al. and 

Ruffner, this volume). Although this period spans 

many human generations, it represents no more than 

several for most oak species, which typically live 150 to 

250 years with some (e.g., white oak) living more than 

600 years (Tirmenstine 1991). Given the longevity of 

oaks and the time that it takes forest stands to develop, 

mature, and be replaced by new growth, managers 

should consider long-term interactions between oaks 

and environmental factors, including fire. The goal of 

this paper is to examine the paleoecology of oak as a 

genus, and to examine the interaction between oaks and 

fire over hundreds and even thousands of years. I start 

from the premise that it is through understanding the 

long-term history of the interaction of fire and oaks that 

we will gain the wisdom required to manage modern 

oak ecosystems and to perpetuate them far into the 

future.

PALEOECOLOGICAL METHODS

It is widely accepted that Native Americans more than 

lightning were the primary source of ignitions that 

burned through eastern forests prior to the arrival 

of Europeans (Bromley 1935; Day 1953; Cronon 

1983). Although opinions differ on the extent of 

Native American burning (Russell 1983), there can 

be little argument that human-caused fires affected 

vegetation development, at least near Native American 

population centers (Patterson and Sassaman 1988). 

The first successful European settlements were in the 

early 17th century in Virginia and Massachusetts, but 

Mann (2002) argued that Europeans influenced Native 

populations, and by inference their effects on vegetation 

through burning, long before that. As early as the late 

15th century with Spanish exploration in the South, and 

perhaps earlier (by Vikings to the north), Europeans 

brought diseases that decimated Native American 

populations throughout the East. To the extent that 

human populations were reduced, the influence of fire 

on vegetation probably was similarly reduced.

There are several methods for exploring fire and 

vegetation history: written accounts by explorers 

and naturalists, dendroecology and fire scar analysis, 

evaluation of 19th century photos in comparison with 

modern photographs (repeat photography), and, in the 

20th century in particular, fire records and quantitative 

descriptions of fire effects. All of these methods have 

been applied effectively to describe vegetation and 

fire interactions before or at the time that Europeans 

settled the West. But few are useful in the East, where 

an understanding of fire regimes and fire-vegetation 

interactions unaffected by Europeans depends on 

a knowledge of how fire, vegetation, and humans 

interacted 500 or more years ago. Trees older than 

about 400 years are rare in the East, especially outside 

of swamps. Historical accounts described the land only 

at the time of observation, and extrapolation to before 

1600 AD is tenuous (Mann 2002). Photographs, written 

records and ecological data from the 19th and 20th

centuries tell us little about what might have occurred 

300 to 400 years earlier.

Thus, paleoecologists have only fossil pollen and charcoal 

analyses of lake and bog sediments with which to 

investigate fire-vegetation interactions for the prehistoric 

period. Fossil pollen analysis, first developed by von Post 

(1916, cited in Davis 1963) in Scandinavia, has been 

used in North America since the 1930’s to reconstruct 

vegetation-climate interactions. Iversen (1941) first used 

charcoal analysis to investigate the effects of Stone Age 

people on vegetation of Denmark. But the technique 

has only recently (since about 1950 initially, and, widely, 

since the 1970’s) been employed in North America 

(Patterson et al. 1987). Both pollen and charcoal analyses 

involve examination of indirect evidence of vegetation 

and fire and are fraught with a series of assumptions and 

constraints. But the data they provide about periods in 

the past for which no other information is available is of 

great value.
	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Fossil Pollen Analysis

Paleoecologists obtain sediments from small ponds, 

lakes, and bogs using coring devices that preserve the 

stratigraphy of the core, with the youngest sediments at 

the top and older ones below. Cores are extracted in 0.5- 

to 1-m lengths and samples are withdrawn at intervals 

that depend on the rate at which sediments are estimated 

to have accumulated and the objectives of the particular 

study. A study of vegetation-climate interactions over 

thousands of years might involve sampling a core at 

intervals of 10 or more cm (100- to 200-year precision). 

Fire-vegetation interactions, are best described by data 

from cores sampled at one to several centimeters (with 

perhaps 5- to 20-year precision).

The theory of pollen analysis is well established. Von 

Post (1916), and, more recently, several North American 

studies observed that pollen is generally well preserved 

in cold, anaerobic, acidic, lake and bog sediments; that 

pollen of different species or genera can be distinguished 

one from another; and that for most species the 

proportion of pollen grains of different types in sediment 

samples approximates the proportion of plants of those 

types on the landscape. Sediments from cores drawn from 

small ponds or wetlands are more likely to reflect local 

conditions (Jacobson and Bradshaw 1981), with cores 

from bogs often containing information about vegetation 

of both the wetland and the surrounding upland.

Not all species can be distinguished based on their 

pollen; this is a particular problem for oaks. None of 

the 36 species native to eastern North America can 

be distinguished one from another based on pollen 

characteristics. Oaks as a group can be distinguished 

from other members of the Fagaceae family, e.g. beech 

and chestnut, and they can be distinguished from the 

hickories, birches, hemlock and pines, so it is largely at 

the level of genera that we can make inferences about fire 

and oak interactions. As with the other wind-pollinated 

species mentioned, oaks produce abundant pollen 

compared to maples and other largely insect-pollinated 

species.

Pollen data usually are expressed as a percentage, with 

the amount of oak pollen, as an example, a percentage 

of all fossil pollen identified on microscope slides 

after sediments are processed to remove inorganic and 

nonpollen organic constituents (Faegri and Iversen 

1989). Percentage data suffer from the inherent 

limitation that if the absolute abundance of one 

constituent (e.g., oak pollen) increases (or decreases) 

dramatically, then the other constituents (e.g., pine, 

hemlock, hickory, birch and whatever other pollen 

types are identified as being present in the sample) must 

decline (or increase), on a precentage basis, even when 

the absolute number of pollen grains remains constant. 

The problem is illustrated with hypothetical data (Table 

1A), where an absolute decline in hemlock pollen is 

reflected in an apparent increase in pine, oak, and other 

types even though they have the same absolute amounts 

in all samples. This problem can be circumvented if a 

known quantity (say 40,000 grains per cubic centimeter 

of sediment being processed) of a marker grain (pollen 

of a type not present in the local flora, e.g., Eucalyptus

pollen for the Eastern United States) is added during 

processing of the sediment and counted along with 

the fossil grains. Where the amount of fossil pollen is 

compared with that of the marker grains, fossil pollen 

can be expressed as an absolute amount of pollen 

(number of grains per unit volume of sediment, see Table 

1B).

If sediment accumulates at the same rate in a basin 

over time, the absolute pollen frequency (APF) allows 

species/genera represented in the data set to be evaluated 

independently of other types (Davis 1965). However, 

sediment accumulation rates are not always the same, 

so APF (number of grains/cm3) is divided by sediment 

accumulation rate (cm/year) to yield a pollen influx 

rate (in number of grains/cm2/year) (Davis and Deevey 

1964). This measure of pollen abundance represents 

an ideal that is not always attainable because sediment 

accumulation rates cannot always be estimated 

accurately.

Cores are dated and sedimentation rates established by 

a variety of methods. Carbon-14 dating can establish 

absolute dates to 30,000 or more years before the 

present. But the accuracy of C-14 dates is on the order 

of decades to 100 years or more (Bradley 1999) and 

declines rapidly in sediments less than about 500 years 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

old. Other isotopes, including Lead-210 (Pb-210), 

are useful for dating younger sediments. Lead-210 is a 

decay product of radon gas, which occurs naturally in 

soils. Concentrations of Lead-210 in sediments are used 

to establish sediment accumulation rates and to date 

sediments that generally are less than 150 to 200 years 

old. Cesium-137, a product of atomic bomb testing in 

the 1940’s, allows dating of younger sediments.

Regionwide changes in the abundance of several pollen 

types have been dated independently. Examples include 

the decline in hemlock (Tsuga canadensis) in eastern 

North America approximately 5,000 years ago (Webb 

1982) and the recent (1900-1920 AD) decline in 

chestnut (Castanea dentata) due to the chestnut blight 

(Anderson 1974). Increases in the abundance of pollen 

of ragweed (Ambrosia spp.) and plantain (Plantago spp.) 

mark the local advent of European agriculture from 300 

to 350 years ago on the East Coast to 100 to 150 years 

ago in Minnesota.

The most precise estimates of sediment accumulation 

rates can be obtained from varved lake sediments, 

i.e. those that have alternating layers of light and 

dark sediment that correspond to changes in seasons 

within a year. A pair of one light and one dark layer 

forms a couplet that indicates a full year of sediment 

accumulation (Fig. 1). Several processes can form banded 

(or laminated) sediments (O’Sullivan 1983), and precise 

chronologies can be established from them. However, 

they are rare, having been identified in no more than a 

dozen or so lakes in Eastern North America.

A

depth Pine # Pine % Oak # Oak %

Hemlock 

#

Hemlock 

% Other # Other % Total #

EXOTIC 

#

500 100 39.2 25 9.8 5 2.0 125 49.0 255 400

503 100 39.2 25 9.8 5 2.0 125 49.0 255 375

506 100 39.2 25 9.8 5 2.0 125 49.0 255 350

509 100 38.5 25 9.6 10 3.8 125 48.1 260 400

512 100 33.3 25 8.3 50 16.7 125 41.7 300 375

515 100 25.0 25 6.3 150 37.5 125 31.3 400 425

B

#/cc #/cc #/cc #/cc #/cc

500 10000 2500 500 12500 25500

503 10667 2667 533 13333 27200

506 11429 2857 571 14286 29143

509 10000 2500 1000 12500 26000

512 10667 2667 5333 13333 32000

515 9412 2353 14118 11765 37647

Table 1.—Hypothetical pollen data showing the effects a large change in the absolute amount of one 

pollen type (hemlock) on the percentages of other types. The number of hemlock grains (A) decreases 

from 150 to 5 from the deepest to the shallowest sample, whereas the number of all other types does not 

change. The decrease in hemlock grains is properly reflected in a sharp decline in hemlock pollen as a 

percentage of the total even though the total number of grains identified declines. But the percentages of 

the total represented by the other three types—pine, oak and all other types combined (other)—increase 

by more than 50 percent. Expressing the pollen as number of grains per cubic centimeter (absolute pollen 

frequency) yields a more appropriate interpretation (B). The addition of 40,000 grains of an exotic pollen 

grain per cc of sediment sample processed, and counting these grains along with the fossil grains, is 

assumed. Expressing data as APF still assumes that each sample represents the same number of years of 

accumulated sediment, an assumption that must be verified by careful dating of many sediment samples 

and the calculation of sediment accumulation rates for different portions of the core being analyzed.
	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

In summary, although imprecise by the standards 

applied to modern ecological studies, fossil pollen 

analysis techniques are largely standardized among 

research labs and limitations are well understood. Pollen 

analysis provides information about time periods for 

which no other information is available and is a useful 

tool for exploring hypotheses about past vegetation 

development and vegetation-environment interactions.

Sedimentary Charcoal Analysis

Sedimentary charcoal analysis is less widely used than 

pollen analysis, and there are a variety of competing 

techniques for quantifying past charcoal production i.e., 

fire activity. The half dozen or so U.S. labs routinely 

performing charcoal analysis tend to use a variety of 

methods that may or may not yield comparable results 

when applied to the same sediments (Patterson et al. 

1987). Recent advances in the theory and methods 

of sedimentary charcoal analysis (Clark 1988a, b) are 

most useful when accurate and precise information on 

sediment accumulation rates is available.

Unlike pollen, which is produced annually in roughly 

the same amounts from similar vegetation types, 

charcoal is produced intermittently from fires that 

occur at different return intervals and that burn with 

different intensities in different fuels across varying 

potions of a landscape. Abundant charcoal is carried 

long distances in towering convection columns from 

large, intense crown fires burning in conifer forests. 

Conversely, charcoal produced by lower intensity 

surface fires burning in deciduous forest (e.g., oak and 

other hardwoods in the Eastern United States) is less 

likely to be transported long distances by air, even when 

large areas burn in a single fire. Short-distance transport 

by air and hydrologic transport in runoff probably 

characterizes these fires. Analysis of pollen data tends 

to focus on running averages that emphasize the yearto-year continuity in pollen production and gradual 

changes in vegetation, whereas fires can elicit dramatic 

changes in charcoal influx from one year to the next, 

and can dramatically affect pollen production within 

the area contributing pollen to a basin.

Patterson et al. (1987) discussed how charcoal is 

produced, transported to deposition sites, and 

eventually preserved in sediments. Larger charcoal 

particles signal locally occurring fires (Clark and 

Patterson 1997; Clark and Royall 1995a), so 

quantifying charcoal by particle size rather than by 

number of fragments has been emphasized. Charcoal 

often is quantified on microscope slides prepared for 

pollen analysis, but Clark (1988b) argued that it is the 

abundance of macroscopic charcoal (i.e., fragments 

longer than 150 to 200 microns) that provides the most 

information about local fire history. This is likely true 

for intense fires burning in heavy fuels in dry conifer 

forests. It is less clear if the generalization applies to fires 

burning through predominantly leaf (nonwoody) litter 

in deciduous forests.

Figure 1.—Photomicrograph of varved sediments from a lake 

in northwestern Minnesota. Thin, horizontal bands of light and 

dark sediments throughout the core are thought to represent 

changes in the chemical composition of sediments related to 

seasonal variations in oxygen content of the bottom waters, 

with each pair of bands representing one year (Foster 1976). 

Broad bands in the middle of the section may have been 

formed by erosion events, and pairs may or may not represent 

one year’s sediment accumulation (Patterson 1978). The core 

section is approximately 10 cm wide by 25 cm long, with the 

youngest sediments at the top of the section.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Charcoal area (the sum of the sizes of all fragments in 

a sample) often is represented relative to fossil pollen 

content (i.e., square microns of charcoal:number of 

fossil pollen grains, or μ2Ch:P) based on the idea that 

local fires produce large charcoal particles while at 

the same time reducing pollen production by killing 

plants. Because fires, and thus charcoal production, 

are transient on the landscape, reconstructing fire 

histories requires sampling sediments at close intervals 

and the analysis of contiguous (or nearly so) analysis of 

samples—what Green and Dolman (1988) refered to 

as fine resolution analyses. It is for this type of analysis 

that varved sediments can be most useful (Swain 1973; 

Clark 1988a). However, they are not required, as shown 

by Motzkin et al.’s (1993) detailed reconstruction of the 

fire and vegetation history of a Cape Cod cedar swamp. 

It is perhaps most important to sample small basins with 

well-defined watersheds relative to the area burned if one 

hopes to identify the response of vegetation to individual 

fires (Patterson and Backman 1988).

THE PALEOECOLOGY OF OAKS

Hundreds of sites have been cored and evaluated for 

fossil pollen content in North America. The National 

Oceanographic and Atmospheric Administration’s 

(NOAA) Fossil and Surface Pollen Data website 

provides access to data for many of these. A map of 

the distribution of sites for which data are archived can 

be viewed at: http://www.ncdc.noaa.gov/paleo/pollen.

html by clicking on the North American portion of the 

Web Mapper box. The cursor can be used to outline the 

eastern portion of the resulting view of North America 

to locate individual sites. Clicking on individual red stars 

allows the viewer to see a pollen diagram for that site, 

or access the original data used to generate the diagram. 

As an example, clicking on the one star in the state of 

Kentucky and then clicking on Diagram (rather than 

Data) provides view of the pollen diagram for Jackson 

Pond (Wilkins et al. 1991). The site has sediments that 

date to at least 20,000 years ago, and the pollen data 

show that oaks have dominated on the surrounding 

upland for approximately 10,000 years.

There are more than 100 sites for Eastern North 

America in the North American Pollen Data Base, and 

diagrams and data are available for each. Data from 

these sites have been merged to provide maps of North 

America which show the abundance of individual pollen 

types at discrete times (generally in increments of 500 

or 1,000 years) in the past. By clicking on the Pollen 

Viewer box, you can access animated reconstructions 

of the spatial and temporal variation in the abundance 

of individual types. Using the scroll box to locate 

Quercus (5,20 and 40%) brings up color-coded range 

and abundance maps for oak for the past 21,000 years. 

Comparisons across many sites of oak pollen abundance 

in modern sediments with the occurrence of oak species 

on the surrounding landscape (as from forest inventory 

data) allow us to make inferences about the importance 

of oak as a component of past vegetation. When oak 

pollen exceeds approximately 5 percent, oak generally 

is at least present within the area contributing pollen to 

the basin. At 20 percent, oak is abundant; at 40 percent 

it dominates the overstory in the surrounding forests 

(Webb 1978; Webb et al. 1978). By clicking on the Play 

button, you can see changes in oak pollen percentages 

over the past 21,000 years and interpret the extent of 

oak trees on the landscape.

Animating the pollen viewer shows that oaks reached 

their current range limit by approximately 10,000 years 

ago and were most important during the Holocene, 

from 9,000 to 10,000 BP. They were the first hardwoods 

to establish and dominate following the retreat of boreal 

spruce-fir forests to the north, and their lower pollen 

percentages after about 9,000 BP is partly a reflection 

of the arrival and establishment of other species of 

the eastern deciduous forest (e.g., beech, the hickories 

and maples and, finally, chestnut). The pollen viewer 

shows a contraction of the area dominated by oaks ( > 

40 percent) during the past 500 years, especially in the 

Central States west of the Appalachians. This probably 

reflects the loss of forest land to agriculture in the 

Midwest. Similar declines (but more recent recoveries) 

in the Northeastern States have been attributed to 

overcutting of oaks for fuelwood in the 18th and 

early 19th centuries (Brugam 1978; Backman 1984). 

Cordwood shortages during this period are documented 

in the historic records for south-coastal New England 

(Stevens 1996).
	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

THE PALEOECOLOGY OF FIRE

The NOAA web site referenced for pollen also provides 

access to fire history data: http://www.ncdc.noaa.gov/

paleo/impd/paleofire.html. This site also links to a Web 

Mapper for North American sites, but the map shows 

only one “charcoal sediment” site east of the Mississippi 

River (at Indiana Dunes National Lakeshore). Pollen 

diagrams with accompanying charcoal data have been 

produced for dozens of sites in the Eastern United States 

by researchers from, among others, the Universities of 

Minnesota, Maine, and Massachusetts; Rutgers, Brown 

and Duke Universities; and the Harvard Forest (Harvard 

University). Several of these sites are included in the 

North American Pollen Data Base, but with no charcoal 

data.

The reasons for the paucity of data for charcoal for the 

NOAA sites are many, but perhaps the most important 

is the lack of a standardized way of reporting charcoal 

results. Summaries of data for some sites have been 

published (Patterson and Backman 1988; Patterson and 

Sassaman 1988; Clark and Royall 1996; Parshall and 

Foster 2002); From these summaries we can generalize 

that just before European settlement, the amount of 

charcoal in sediments decreased from west to east, with 

the lowest values in the mountainous regions of New 

England (Patterson and Backman 1988). For the most 

part, sedimentary charcoal content decreased with the 

arrival of European settlement west of the Appalachians, 

but increased in New England. This pattern generally is 

consistent with perceptions of the role of fire in eastern 

North America before and after European settlement 

(Pyne 1982).

THE PALEOECOLOGY OF OAK

AND FIRE

Access to charcoal data in the same NOAA formats 

available for pollen would greatly facilitate an evaluation 

of the prehistoric role of fire in maintaining oak forests. 

For now, researchers are left with the evaluation of 

fire-oak interactions on a site-by-site basis. Reviews are 

available but the results are ambiguous. Clark and Royal 

(1995) documented a shift from northern hardwoods to 

white pine/oak forests with the establishment of a local 

Indian settlement about 1450 AD in southern Ontario, 

at Crawford Lake, and attribute the shift to Indianignited fires. In support of the role of Native American 

burning in the maintenance of oak forests, Delcourt and 

Delcourt (1997) used pollen, charcoal, and archeological 

data to conclude that “Native Americans played an 

important role in determining the composition of 

southern Appalachian vegetation through use of fire.” 

They argued that fires would have stimulated sprouting 

“thereby increasing the abundance of chestnut and oaks 

growing in open groves.”

Campbell and McAndrews (1995) disagreed with Clark 

and Royall’s (1995b) Crawford Lake interpretation 

despite an apparent sharp increase in sedimentary 

charcoal at about the time that oak and pine percentages 

increased at the expense of elm, sugar maple, and beech. 

They attributed the change in forest composition to 

climate anomalies associated with the Little Ice Age. 

Clark (1995) defended the fire hypothesis but careful 

examination of the pollen and charcoal data (Figure 

3 in Clark and Royall 1995b) apparently shows that 

oak pollen percentages began to increase just before 

(ca. 1400 AD) a major fire, as indicated by the highest 

charcoal influx values (centered on ca. 1450) in the core. 

Oaks producing pollen that contributed to the increase 

in percentages ca. 1400 AD probably established as early 

as 1350, given the several decades that it takes oaks to 

produce pollen after establishing from seed. Peak pollen 

values for oak (30 to 40 percent) are in sediments that 

postdate the ca. 1450 fire (approximately from 1500 to 

1800). Clark (1995) notes that the Crawford lake site 

appears to show a different set of transitions (with a 

different cause—Native American burning vs. climate 

change?) compared to other sites in the region. At 

Devel’s Bathtub in west-central New York, Clark and 

Royal (1996) found abundant oak pollen throughout 

the Holocene, but surprisingly little charcoal in the 

sediments. On the basis of data from this site, they 

questioned the dependence of oak on fire “at Devil’s 

Bathtub and other sites in the northeast” (Abrams 

2002).

Following the publication of their discussion with 

Campbell and McAndrews regarding the interpretation 

of the Crawford Lake site, Clark and Royall (1996) 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

reviewed local and regional sediment charcoal for 

evidence of fire in presettelment northeastern North 

America. Thye reported that “We did not find evidence 

for fire in mixed oak forests, where it has been speculated 

that fire might be necessary for oak recruitment.” 

Abrams (2002), reviewed several published pollen 

and charcoal diagrams, including those for Crawford 

Lake and Devil’s Bathtub. Despite acknowledging the 

reservations of Clark and Royall (1996), he concluded 

that “many paleoecological papers show an intrasite 

increase in regional or local charcoal with increases in 

oak abundance.”

My own review of published and unpublished work 

for this paper supports the somewhat ambiguous 

interpretation suggested by Clark and Royall. It is clear 

that at many sites charcoal is abundant when oak is 

dominant, or that across sites, samples from sites with 

more oak tend to have more charcoal than those from 

sites surrounded by more mesic, less fire-prone forest 

types. Crawford Lake notwithstanding, what appears 

to be lacking is evidence that an increase in fire has 

caused oak to become more abundant where it was less 

so. Most fire managers would acknowledge that litter 

fuels in oak forests are more prone to burn than those in 

mesic forests. Indeed, the northern hardwood forests of 

western New England are widely referred to by foresters 

as “asbestos forests” (Borman and Likens 1979), an 

acknowledgement of the low flammability of beech, 

maple, and birch litter. But questions remain: Is there 

more oak pollen in sediment samples with abundant 

charcoal because fire favors oak-dominated vegetation? Is 

there more charcoal in sediments with abundant charcoal 

because oak fuels are more likely to burn?

I believe that insights can be gained by examining 

individual sites. I next use data from sites in Virginia, 

southern New England/Long Island, and on the coast of 

Maine to illustrate some of what we have learned about 

the interaction between fire and oaks along the Atlantic 

Seaboard. Following the example of Jacobson (1979), I 

have chosen pairs of sites in each of the three regions to 

illustrate how oaks growing on different sites respond 

differently to fire.

Western Virginia

We reconstructed fire and vegetation histories for 

two different sites near Staunton in western Virginia 

(Patterson and Stevens 1995). Brown’s Pond, at an 

elevation of 2,000 feet in Bath County (38°08 ́50′′ N, 

79°36 ́23′′ W), lies in a protected cove and is surrounded 

by mixed mesophytic forests dominated by red and white 

oaks and a variety of associated hardwood species. Pines 

occur only occasionally on the surrounding uplands. 

Soils are deep (greater than 60 inches to bedrock) loams. 

Pollen (but not charcoal) data for sediments dating to 

17,350 BP are available in the North American Pollen 

Data Base (Kneller and Peteet 1993). We obtained a 

90-cm-long core from the site in 1994 and analyzed 

the sediments for pollen and charcoal content (Fig. 2). 

Increases in pollen of agricultural indicators (chiefly 

ragweed) mark the advent of cultivation by Europeans 

300 years ago. Oaks have been the dominant species in 

the Brown’s Pond watershed, with pollen ranging from 

40 to 60 percent (of all pollen grains identified) in all 

samples. Charcoal to pollen ratios (Ch:P) range from 0 

to more than 400. Our work on Mount Desert Island, 

Maine suggests that Ch:P values in excess of 150 to 200 

indicate fires occurring in the watershed of small ponds, 

whereas lower values probably represent charcoal blown 

in from afar (Patterson and Backman 1988; Clark and 

Patterson 1997). Although individual peaks in charcoal 

abundance are evident, the sampling interval was 

broad and identifying individual fires and estimating 

the interval between them are not possible. Still, we 

concluded that fires have been part of the local landscape 

throughout the 6,000-year period represented by the 

core. Charcoal values were somewhat higher before 

Europeans arrived, but the overall abundance of fire on 

the landscape did not change with their arrival.

Green Pond is approximately 30 miles southeast of 

Brown’s Pond at an elevation of 3,200 feet in Augusta 

County (37°56 ́23′′ N, 79°02 ́51′′ W). It lies on 

an exposed ridge known locally as Big Levels and is 

surrounded by chestnut (Quercus prinus) and black (Q. 

velutina) oaks, and pitch pine (Pinus rigida) over dense 

thickets of ericaceous shrubs. Soils are very stony loamy 

sands with depths to bedrock of 40 to 48 inches. A 
1	 Fire 0 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

35-cm-long sediment core obtained in 1994 represents 

approximately the last 900 years. Oaks (40 percent of 

total pollen) dominated the uplands prior to European 

settlement about 250 years ago. Pines, which like oaks, 

produce abundant pollen, were less important, and 

charcoal values were somewhat lower (Ch:P = 0-300) 

than observed at Brown’s Pond. One or more major 

fires apparently burned near the pond about 150 to 250 

years ago, with charcoal abundance increasing to 400 

to 600 before declining in recent sediments (Fig. 3). 

The increase in fire activity was followed by an increase 

in pine and a decline in oak. Although oaks remain an 

important component of the modern vegetation, recent 

pine pollen percentages are more than double those prior 

to settlement.

Results from Brown’s and Green ponds suggest that 

on mesic soils in protected locations of the southern 

Appalachians, light surface fires may contribute to 

maintaining oaks as a dominant component of mixed 

mesophytic forests—perhaps by selecting against more 

shade tolerant but less fire tolerant mesophytic hardwood 

species. On exposed sites with poorer soils where shadetolerant (mesic-site) species compete less well (e.g., 

at Green Pond), oaks may persist with less fire. More 

frequent fires, at least some of which are likely to be 

more severe on shallow soils, may favor the establishment 

of pines and increase their importance on the landscape. 

However, longer lived oaks still persist, even in the face 

of competition from dense shrub understories, which 

may prevent pines from regenerating without fire 

(Patterson and Stevens 1995).

South Coastal New England and Long Island

Deep Pond lies at an elevation of 28 feet on the 

Ronkonkama Moraine in northeastern Suffolk County, 

Long Island, New York (40° 56 ́ 11′′N, 72° 49 ́ 52′′W). 

The uplands surrounding the pond are dominated by 

black and white oak and pitch pine. Soils are deep, 

loamy sands. A sediment core spanning the past 2,200 

0

5

10

15

20

25

30

35

40

45

50

55

60

65

70

75

80

85

90

DEPTH IN SEDIMENT (cm)

ca. 1700 AD

650 ± 60

4210 ± 40

C-14 AGE (BP)

20

%

0

PINE

20 40 60

%

0

OAK

%

0

GRASSES

20

%

0

AGRICULTURAL INDICATORS

200 400

microns^2 : upland pollen

0

CHARCOAL

BROWN'S POND

Bath County, Virginia

FOSSIL POLLEN AND CHARCOAL

1994

Analyst: A. Stevens

Figure 2.—Pollen and charcoal profiles for 

Brown’s Pond, Bath County western Virginia. 

The date of local settlement by Europeans 

(dashed line) is from historical accounts.

BROWN'S POND

Bath County, Virginia

FOSSIL POLLEN AND CHARCOAL

1994
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 11

years was recovered in 1981 (Backman 1984). Oaks 

dominated throughout the sample period, though 

percentages have declined somewhat, with pitch pine 

increasing in importance since European settlement 

about 300 years ago (Fig. 4). Charcoal values are high, 

averaging 400 to 600 before settlement and increasing to 

500 to more than 1,000 in recent centuries. As at Green 

Pond in Virginia, recent increases in fire activity appear 

to favor pitch pine.

Charge Pond, at an elevation of 66 feet in Plymouth 

County, Massachusetts (41° 48 ́ 58′′N, 70° 40 ́ 31′′W), 

occupies a kettle hole in outwash of the Wisconsin 

glacial stage. Course, sandy soils currently support a 

mixture of pitch pine and scrub oaks. The area around 

the pond burned most recently in a catastrophic fire in 

1957. Backman (1985) described the fire and vegetation 

history for the past several hundred years based on his 

analysis of a core obtained in 1981 (Fig. 5). Fire activity 

has been high throughout the last several hundred years, 

as evidenced by charcoal values which rise from 500 to 

1,000 before European settlement to 4,000 to 5,000 

during the post-settlement period. Oaks and white pine 

(Pinus strobus) were dominant before settlement. Because 

tree oaks are more likely to occur with white pine, we 

assume that white and black oak were more important 

than scrub oaks (Patterson and Backman 1988). Today, 

the vegetation on the upland is referred to as pitch 

pine-scrub oak barrens, but our data suggest that these 

species—especially pitch pine—have been dominant 

only since fire activity increased following European 

settlement.

Data from Deep and Charge ponds suggest that on 

course-textured soils, where pines are better able to 

compete with oaks, oaks survive at moderately high 

fire activity, but composition has shifted towards an 

increasing importance of pitch pine with the high fire 

activity during the post-settlement period. A more 

detailed examination of the pollen data suggests that 

with increasing fire activity, more mesic hardwood 

species, including hickory (Carya spp.), maple and 

beech, decline in importance.

Figure 3.—Pollen and charcoal profiles for 

Green Pond, Augusta County, western Virginia. 

The date of local settlement by Europeans 

(dashed line) is from historical accounts.

0

5

10

15

20

25

30

35

DEPTH IN SEDIMENT (cm)

ca. 1750 AD

730 ± 60

C-14 AGE (BP)

20 40 60

%

0

PINE

20 40

%

0

OAK

%

0

GRASSES

20

%

0

AGRICULTURAL INDICATORS

200 400 600 800

microns^2 : pollen

0

CHARCOAL

GREEN POND

Augusta County, Virginia

FOSSIL POLLEN AND CHARCOAL

1994

analyst: A Stevens

GREEN POND

Augusta County, Virginia

FOSSIL POLLEN AND CHARCOAL

1994
12	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Coastal Maine

Mount Desert Island in Hancock County, Maine, is 

a 120,000-acre mountainous, ocean island 40 miles 

south of Bangor. Diagrams for two small ponds provide 

insights about the interaction between fire and oaks 

near their northern range limit. Only red and bear oak 

(Quercus ilicifolia) occur naturally on the island today.

Sargent Mountain Pond is at an elevation of 1,145 

feet (44° 20 ́ 04′′N, 68° 16 ́ 10′′W). Mature spruce 

(Picea spp.) forests on shallow-to-bedrock, granitic 

soils surround the small mountain pond, which lies 

just outside the western boundary of the great Bar 

Harbor Fire, which burned nearly 20,000 acres on the 

Island in October 1947. Oaks are not present in the 

pond’s small watershed but there is a stand of red oak 

regenerated by the 1947 fire about 1 mile to the north. 

Pollen and charcoal analysis of a 30-cm-long sediment 

core provides information on the fire and vegetation 

history of the watershed for the past 400 years (Fig. 6). 

At least two fires burned the surrounding upland, one 

about 1700 AD, 60 years before the first permanent 

settlement on the Island at Somesville 2 miles to the 

west, and a second in the mid-19th century. The second 

fire probably occurred in conjunction with the logging 

of the mountain’s virgin stands of spruce. Charcoal at 

background levels of less than 200 μ2

 forthe rest of the 

core shows that no other fires have burned within the 

watershed during the period of record. The two discrete 

fires separated by 150 years allow interpretation of postfire succession—alder (Alnus spp.), followed by birch, 

and finally spruce—that is consistent with what has 

been observed following early and mid-20th century fires 

(Patterson et al. 1983; Patterson and Backman 1988). 

Oak pollen occurs at generally less than 5 percent and 

rises only slightly following fires, perhaps reflecting the 

increased importance of regional pollen following standreplacement fires on the watershed.

Figure 4.—Pollen and charcoal profiles for 

Deep Pond, Suffolk County, Long Island, New 

York. The date of local settlement by Europeans 

(dashed line) is from historical accounts.

0

50

100

150

200

250

300

350

400

450

500

550

600

650

700

750

800

850

900

950

1000

1050

1100

1150

1200

1250

1300

1350

1400

1450

DEPTH IN SEDIMENT (mm)

ca. 1680 AD

2180 ± 130

C-14 AGE (BP)

20

%

0

PINE

20 40 60

%

0

OAK

%

0

GRASSES

20

%

0

AGRICULTURAL INDICATORS

1000 2000

microns^2 : upland pollen

0

CHARCOAL

DEEP POND

Wadding River, Suffolk Co., LI, NY

FOSSIL POLLEN AND CHARCOAL

1981

analyst: A.E. Backman

DEEP POND

Wadding River, Suffolk Co., LI, NY

FOSSIL POLLEN AND CHARCOAL

1981
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 13

Lake Wood, at a lower elevation (35 feet) and 6 miles 

northeast of Sargent Mountain Pond (44° 24 ́ 27′′N, 68° 

16 ́ 06′′W), lies in a glacially scoured valley with soils 

derived from course glacial debris overlain in places by 

marine clays. The entire watershed of Lake Wood burned 

in the 1947 fire. The vegetation today is dominated by 

paper (Betula papyrifera) and gray birch (B. populifolia), 

red (Pinus resinosa) and white pine, and occasional red 

oaks. A small stand of hemlock in the southeastern part 

of the watershed regenerated following the 1947 fire. 

Hemlock is uncommon in the forests of Mount Desert 

Island.

Like Brown’s Pond in western Virginia, we have a long 

record of fire-vegetation interactions from Lake Wood. 

Unlike the Virginia core, the Lake Wood core was 

sampled and analyzed with very fine resolution; sampleto-sample precision is on the order of 20 to 40 years. 

The pollen data suggest oak has been present as at least 

a minor component of the local vegetation for much 

of the past 6,000 years despite substantial changes in 

species dominance. Hemlock, white pine, birch, sugar 

maple, and beech (not shown in Fig. 7) dominated until 

about 2,000 years ago when spruce (plus cedar and fir) 

replaced the hemlock and northern hardwoods. The 

charcoal record clearly documents the local occurrence 

of the 1947 Bar Harbor fire at the top of the profile. 

This fire occurred during the driest month in more 

than 150 years of recorded weather history in Maine 

(Baron et al. 1980), and may have been the most severe 

fire to occur on the watershed during the entire 6,000 

year period represented by the core. However, different 

fire regimes appear to dominate at different times, and 

the abundance of oak varies with them. Approximately 

3,000 to 6,200 BP as many as five or six charcoal peaks 

occurred, usually in conjunction with changes in the 

abundance of hemlock. We are investigating whether 

these fires followed declines in hemlock caused by 

insects, disease, or blowdown, or themselves were the 

cause of the declines (unpublished data), but oaks are 

Figure 5.—Pollen and charcoal profiles for 

Charge Pond, Plymouth County, Massachusetts. 

The date of local settlement by Europeans 

(dashed line) is from historical accounts.

0

5

10

15

20

25

30

35

40

DEPTH IN SEDIMENT (cm)

ca. 1800 AD

20 40

%

0

WHITE PINE

20

%

0

PITCH PINE

20 40

%

0

OAK

%

0

GRASSES

20

%

0

AGRICULTURAL INDICATORS

2500 5000

microns^2 : pollen

0

CHARCOAL

CHARGE POND

Plymouth County, MA

FOSSIL POLLEN AND CHARCOAL

1981

analyst: A.E. Backman

CHARGE POND

Plymouth County, MA

FOSSIL POLLEN AND CHARCOAL

1981
1	 Fire 4 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

clearly more abundant during this period than during 

the subsequent 1,000-year period (2,000 to 3,000 BP) 

with no fires. The shift from hemlock to spruce in the 

region at 2,000 BP is accompanied by an overall increase 

in fire activity, but oak does not increase again until well 

after this change in vegetation (and fire regime). Oak 

pollen percentages at the surface of the core are higher 

(at 11 percent) than at any time since about 3,700 BP.

In conjunction with a companion site (The Bowl) on the 

Island for which we have comparable detail for an even 

longer period of time (unpublished data), the detailed 

sampling of the Lake Wood core over a long period 

provides us, with the opportunity to seek answers to 

the questions posed earlier: Do changes in fire regimes 

cause changes in oak abundance? Do the increased 

flammability of oak fuels cause more frequent fires on 

the landscape? Our analyses to answer these questions 

are incomplete but the evidence for this one site where 

fire has historically been infrequent on the landscape 

suggests at least a correlation between fire activity and 

oak abundance.

SUMMARY AND CONCLUSIONS

Paleoecological evidence, such as the fossil pollen and 

charcoal data presented here, is incomplete, fraught 

with uncertainties, and dependent on assumptions that 

often are difficult to evaluate. But they are the only 

data available that allow us to make inferences about 

fire and vegetation before the time of recorded history. 

There is value in considering the long-term interactions 

between fire and vegetation, especially for long-lived 

forest trees like the eastern oaks, because processes like 

fire, which influence stand initiation and development, 

have secondary effects that are realized over decades 

or even centuries. Changes in fire return intervals, the 

interaction between fire occurrence (and effects) and 

climate (which itself changes), and the rare occurrence 

(in some systems) of catastrophic fires, must be studied 

over periods far longer than fire scientists are accustomed 

to contemplating.

In this paper I have examined the value of using 

paleoecological data to investigate the response of oaks 

as a group to fire. Chief among the difficulties is that 

Figure 6.—Pollen and charcoal profiles for 

Sargent Mountain Pond, Hancock County, 

Maine. The date of local settlement by 

Europeans (dashed line) is from historical 

accounts and Lead-210 analysis.

0

50

100

150

200

250

300

DEPTH IN SEDIMENT (mm)

ca. 1760 AD

ca. 1700 AD

20

%

0

SPRUCE

20

%

0

PINE

20 40

%

0

BIRCH

%

0

OAK

20 40

%

0

ALDER

%

0

GRASSES

20

%

0

AGRICULTURAL INDICATORS

200 400 600

microns^2 : upland pollen

0

CHARCOAL

SARGENT MOUNTAIN POND

Mt. Desert Island, Maine

FOSSIL POLLEN AND CHARCOAL

1980

Analyst: A.E. Backman

SARGENT MOUNTAIN POND

Mt. Desert Island, Maine

FOSSIL POLLEN AND CHARCOAL

1980
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 15

we cannot distinguish among any of the dozens of oak 

species based on their pollen morphology. Similarly, we 

often can only guess at how changes in the abundance of 

charcoal in sediments relate to the intensity, severity, or 

extent of individual fires. And it is only under unusual 

circumstances that we can identify individual fires and 

their effects on vegetation using fossil data. The detailed 

analyses for the Sargent Mountain Pond and Lake Wood 

sediments are unique in this regard.

Yet fossil data do provide information that may be 

useful in the future management of oak forests. Chief 

among the lessons we have learned is that it is difficult to 

generalize about the interaction between fire and oaks on 

the centuries-long time scales that are relevant to forest 

stand development and change. Oaks have persisted 

for thousands of years in a fire regime apparently 

characterized by relatively infrequent, low-intensity fires 

at Brown’s Pond in western Virginia. But oaks also have 

persisted for millennia under frequent, probably higher 

intensity surface fires at Deep Pond on Long Island. 

At Green Pond, surrounded by more xeric soils and 

vegetation at higher elevations in Virginia, one or more 

severe fires at the time that Europeans arrived changed 

the balance between oaks and pines. The same is true for 

Charge Pond in southeastern Massachusetts. In neither 

case were oaks reduced to minor components of the 

vegetation, but the balance between oak and pine was 

clearly shifted toward the later. On the Maine coast, near 

the northern limits of the range of oaks, fire at times has 

been rare, and oaks have been a minor component of 

the vegetation as a whole. Yet when fire return intervals 

Figure 7.—Pollen and charcoal profiles for 

Lake Wood, Hancock County, Maine. The date 

of local settlement by Europeans (dashed line) 

is from historical accounts.

0

50

100

150

200

250

300

350

400

450

500

550

600

DEPTH IN SEDIMENT (cm)

6240 ± 110

5960 ± 120

4930 ± 90

3830 ± 100

3090 ± 90

2370 ± 100

1970 ± 80

ca. 1760 AD

C-14 AGE (BP)

20

%

0

SPRUCE

20 40

%

0

HEMLOCK

20 40

%

0

WHITE PINE

20 40

%

0

BIRCH

20

%

0

OAK

20

%

0

ALDER

%

0

GRASSES

%

0

AGRICULTURAL INDICATORS

200 400 600

microns^2 : pollen

0

CHARCOAL

LAKE WOOD

Mount Desert Island, Maine

FOSSIL POLLEN AND CHARCOAL

1990

analyst: N.E.R. Drake

SARGENT MOUNTAIN POND

Mt. Desert Island, Maine

FOSSIL POLLEN AND CHARCOAL

1980
1	 Fire 6 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

are reduced (i.e. fires occur more often), oaks seem to 

compete better with more shade-tolerant conifers and 

hardwoods.

In the Upper Midwest, Jacobson (1979) found that 

white pine migrated into oak forests, perhaps in 

response to a “slight decrease in fire frequency” at various 

times during the Holocene. This finding is counter 

to those for the Green and Charge ponds in the East. 

Jacobson did not provide charcoal data to support his 

hypothesis, but it is likely that fires were so frequent in 

the oak savannah stands he postulates that pines would 

not have been able to survive. White (1983) found that 

annual burning maintains oak savannas with grassy 

understories in prairie-forest transition stands in eastcentral Minnesota. Jacobson assumed that white pine 

cannot tolerate return intervals of less than 20 years.

There is little evidence from sites on the East Coast 

that fires were frequent enough to give rise to savannas 

like those of the Upper Midwest. At none of the sites 

I have discussed has an increase in charcoal, either 

before or after European settlement, coincided with 

a substantial increase in grass pollen. Stevens (1996) 

suggested that frequent Native American burning on the 

island of Martha’s Vineyard, Massachusetts, may have 

been responsible for the coincidence of high charcoal 

values and high oak and grass pollen percentages. She 

also found abundant charcoal with abundant oak and 

grass pollen in post-settlement sediments, but it may 

have been grazing and cultivation as much as fire that 

was responsible for this correlation, which was largely 

restricted to coastal sites regionally (Foster and Motzkin 

2003).

Recent work at the Harvard Forest emphasized the 

variety of opinions on the importance of interactions 

between fire and oaks in the Northeast. Parshall and 

Foster (2002) generally discountrd the importance of 

fire relative to climate and soil factors as influencing 

regional variation in the vegetation of the oak region 

of southern New England. But Foster et al. (2002) 

postulated that fire is important, albeit at lower levels 

than along the coast, in maintaining oak and chestnut 

forests in central Massachusetts. Their work shows that 

detailed reconstructions of fire and vegetation histories 

at selected sites hold the promise of further advances 

in our understanding of the long-term relationship 

between fire and oaks. However, our interpretation of 

historical data must be informed by a knowledge of 

fire-vegetation interactions from contemporary studies. 

Long-term prescribed burning experiments, like those 

conducted for nearly half a century at the Tall Timbers 

Research Station in Florida (Stoddard 1962), will 

increasingly provide information that will allow us to 

better interpret the paleoecological record. 

ACKNOWLEDGMENTS

Thanks go to Kennedy Clark, Matthew Dickinson, 

and an anonymous reviewer for useful comments on 

an earlier version of this paper. Several former students, 

including Andrew Backman, Anne Marie Hine, Natalie 

E.R. Drake, and Andrea Stevens, analyzed pollen and 

charcoal in sediments generating data that contributed 

to diagrams published here.

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2	 Fire 0 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Fire scars reveal variability and dynamics of eastern

fire regimes

Richard P. Guyette1†, Daniel C. Dey2, Michael C. Stambaugh1, and Rose-Marie Muzika1

Abstract.—Fire scar evidence in eastern North America is sparse and complex but shows promise in 

defining the dynamics of these fire regimes and their influence on ecosystems. We review fire scar data, 

methods, and limitations, and use this information to identify and examine the factors influencing 

fire regimes. Fire scar data from studies at more than 40 sites in Eastern North America document fire 

regimes in forests with oak. Fire frequency was highly variable in both time and space even at regional 

scales (less than 500,000 ha). Many sites burned frequently (2- to 3-year mean fire intervals) while 

nearby sites (less than 40 km distant) burned infrequently (mean fire intervals more than 20 years). The 

fire scar record shows that major factors controlling temporal differences in fire regimes are changes in 

human population density, culture, and annual drought. Spatial differences in fire regimes are influenced 

by regional temperature, human population density, and topographic resistance to the spread of fire. 

Severe fire years (more than 10 percent of trees scarred at the sites) were associated with strong regional 

droughts that covered most of the Eastern United States and southern Ontario, Canada. Major fire years 

in Eastern North America occurred about 3.6 times per century before suppression efforts in forests with 

an oak component. Fire regimes with numerous human ignitions were more influenced by droughts. 

We synthesize mean fire intervals during the pre-European settlement period using an empirically 

derived regression model. The model was developed using two variables to predict broad scale spatial 

differences in fire frequency based on fire interval data derived from dendrochronologically dated fire 

scarred trees. Sixty-three percent of the variance in mean fire intervals was explained by mean maximum 

temperature and 12 percent by mapped human population density and historical documentation. 

The model is used to map coarse scale fire intervals in forested regions of the Eastern United States.

fire locations, and drought. We examine fire history 

in forests of Eastern North America during the last 

three centuries, emphasizing forests that have an oak 

(Quercus spp.) component. We document and analyze 

fire regimes using dated fire scars and examine variables 

that are hypothesized to influence these fire regimes. 

Fire scar data from more than 60 sites in 16 states and 

the province of Ontario (Table 1) are used to document 

fire regimes and examine their temporal and spatial 

variability. To present a more balanced regional analysis 

and overview, we excluded a considerable amount of data 

from Missouri (more than 20 fire history sites).

Dendrochronological analysis of fire scars and the 

reconstruction of fire history from scar data have many 

positive and precise features as well as limitations. 

Fire scars on oaks provide a detailed record of fire 

events, such as exact locations and dates (Guyette and 

Stambaugh 2004; Smith and Sutherland 1999). Each fire 

scar potentially has a spatial resolution of less than 1 m 

and a temporal resolution of less than one year. These 

characteristics generally set this method apart from other 

1

School of Natural Resources, University of Missouri, 

Columbia, MO 65211; 2USDA Forest Service, Northern 

Research Station, Columbia, MO 65211; 

†

Corresponding 

author, 573-882-7741, email: guyetter@missouri.edu.

INTRODUCTION

Wildland fire is an important disturbance influencing 

oak forests (Johnson et al. 2002; Wade et al. 2000). 

Changing forest composition and fire risk have given 

the history of wildland fire regimes new relevance. 

Owing to the longevity of trees, forest composition often 

changes slowly in response to fire and at time scales 

well beyond the lifespan of single human observers. 

To understand how and why forests are changing, 

we often can benefit from a longer-term perspective. 

Dendrochronology can provide this perspective with 

high resolution spatio-temporal data for periods of more 

than 300 years (several generations of trees) in Eastern 

North America. This period spans changes in human 

societies from subsistence to fire suppression. Tree-ring 

dating of fire scars and climate reconstructions from the 

width of tree rings provide long records of fire dates, 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 21

Site name State Forest 

Type

Trees and forest type Pre-Euro-fire 

interval 

Post- Eurofire intervalSite data reference

Green Mts. VT NH red pine forest na 18 Engstrom and Mann 1991

Basin Lake ON NH pine-oak woodland 21 na Guyette and Dey 1995a

Papineau L. ON NH oak-pine forest 29 7 Dey and Guyette 2000

Opeongo ON NH pine-hardwoods 29 25 Guyette and Dey 1995b

Bracebridge ON NH red oak forest 11 na Guyette and Dey 1995c

Joko ON NH pine hardwoods 13 na Dey and Guyette 2000

Seagan ON NH pine hardwoods 17 na Dey and Guyette 2000

Sault Ste. Marie ON NH pine hardwoods 26 73 Alexander et al. 1979

Big Bay MI NH pine- hardwoods 25 na Torretti 2003

Itasca SP MN NH conifer hardwood 25 36 Spurr 1954

Itasca SP MN NH conifer-hardwood 29 13 Clark 1990

Itasca SP MN NH conifer-hardwoods 30 9 Frissell 1973

Costal sand MI NH costal pine forests 18 88 Loope and Anderton 1998

Upland sand MI NH upland pine forests 23 29 Loope and Anderton 1998

Savage Mt. MD CH oak forest 8 8 Shumway et al. 2001

Oreton OH CH white oak forest na 4 Sutherland 1997

Pike Knob WV CH white oak forest na 14 Schuler and McClain 2003

Lemm Swamp TN CH oak woodlands 5 8 Guyette and Stambaugh 2005

Saltwell TN CH oak woodlands 6 8 Guyette and Stambaugh 2005

Richland Creek TN CH oak woodlands 5 4 Guyette and Stambaugh 2005

Huckleberry TN CH oak woodlands 6 4 Guyette and Stambaugh 2005

Boone Barrens IN CH oak savanna 19 5 Guyette et al. 2003

Brush Mt. VA CH pine – hardwood na 8 Sutherland et al. 1992

Mill Hollow MO CH oak-pine forest 6 2 Guyette and Cutter 1997

MOFEP 4 MO CH oak –pine forest 3 na Guyette and Stambaugh 2004

MOFEP 5 MO CH oak –pine forest 4 na Guyette and Stambaugh 2004

Mahans Crk. MO CH oak –pine forest 3 4 Guyette and Cutter 1997

Mill Creek MO CH oak-pine forest 4 4 Guyette and Cutter 1997

Hartshorn MO CH oak-pine woodland 3 2 Guyette and Cutter 1997

Shannondale MO CH oak forest 13 4 Guyette and Cutter 1997

Gee Creek AR CH oak-pine forest 15 2 Guyette and Spetich 2003

Gobblers Knob AR CH oak-pine forest 16 2 Guyette and Spetich 2003

Vermillion MO CH pine-oak forest 3 na Guyette 1997

Rd1645 MO PT oak woodland 4 8 Cutter and Guyette 1994

Pleasant Prairie WI PT oak savanna na 7 Wolf 2004

Ava Glades MO PT redcedar-oak glades 6 11 Guyette and McGinnes 1982

White Ran. MO PT oak woodland 4 8 Dey et al. 2004

Caney Mt MO PT oak savanna 5 7 Guyette andCutter 1991

Loess Hills MO PT bur oak woodland 8 4 Guyette and Stambaugh 2005

Granny Gap AR SP pine–oak forest 5 4 Guyette and Spetich 2003

Chigger Rd AR SP pine-oak forest 13 2 Guyette and Spetich 2003

GSMNP TN SP pine – oak forests na 13 Harmon 1982

Table 1.—Mean fire intervals from fire scar studies in eastern North America (NH = northern hardwoods, 

CH = Central Hardwood, SP = Southern Pine-Hardwoods, PT = Prairie-Forest Transition; na = not available)
22	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

fire history methods (Table 2). However, a number of 

problems may arise when developing and interpreting 

the fire scar record. A woody growth increment without 

a scar is only weak evidence of “no fire” because many 

trees are resistant to scarring. In Eastern North America, 

challenges often arise when constructing fire scar 

histories, such as the limited availability of preserved 

fire scars, limited abundance of closely spaced recorder 

trees, differences in record length, and sampling methods 

that are defined by the limits of the resource. However, 

even studies with the most severe sampling problems rise 

above the alternative: an absence in our knowledge of 

fire history. Problems in developing fire history from fire 

scars (and how we address them in this paper) include:

1. Problem: Many trees have thick bark and are not 

scarred in fires. Thus, the lack of a scar does not 

mean the absence of fire. Solution: Minimize the 

use of studies based only on a single or several trees.

2. Problem: Stand replacement fires are not 

included in fire scar studies because all fire scars 

occur on survivor trees. Solution: Infer from the 

number of trees scarred when larger and more 

severe fires occurred.

3. Problem: Tree species differ with respect to 

recording fire history. This can result in errors 

when comparing histories. Solution: Sample a 

sufficient number of trees to develop a composite 

fire interval that reflects the actual frequency of fire 

in an area.

4. Problem: Spatial extent of a study site influences 

the inferred fire frequency. Solution: Confine 

analysis to sites of several hundred hectares (less 

than 4 km2)

5. Problem: The number of sample trees in any 

given year may affect the number of fires detected. 

Solution: Truncate data from early periods when 

there were few recorder trees in the record.

6. Problem: Many fire scar chronologies have long, 

open-ended intervals that cannot be interpreted 

as true intervals. Solution: Estimate mean fire 

intervals as years per fire.

The goals of this review and synthesis are to examine the 

long-term and large-scale factors that affect the spatial 

and temporal variability of fire regimes in Eastern North 

America. Specific objectives are to examine climate, 

Table 2.—Methods of documenting fire history, their data acquisition and characteristics

Method Sources Resolution Time depth Limitations and advantages

Dendrochronology Crossdated fire 

scars from trees, 

remnant wood. 

Also ring widths, 

stand age

Seasonal to 

annual with high 

precision

Commonly 

two to four 

centuries, rarely 

more

Stand replacement fires 

not well documented, high 

resolution in time and space 

GLO notes Historical archives None About 150 to 

200 years ago

Inferential, spatially explicit 

information 

Charcoal sediment Lakes, peat Variable, decadal 

to centuries

Centuries to 

12,000+ years 

Local sources and low 

resolution, long-term data

Written accounts Historical 

literature and 

travel accounts

Low to daily in 

journals

About 500 years Nonquantitative, often 

biased, includes cultural 

information

Alluvial charcoal Alluvial sediments About 500 years 

or the limits of 

carbon dating

1000 to 

20,000+ years

Low resolution, record of 

large, intense, sedimentproducing fires
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 23

human, and landscape influences on fire regimes using 

data from existing fire scar histories in the eastern oak 

forests. We present a preliminary synthesis via regression 

modeling, using the effects of temperature and human 

population on fire regimes.

Review

Numerous studies (Table 1) have used fire scar data 

to reconstruct fire history from the four major oak 

ecoregions and forest types of Eastern North America 

(Bailey 1997; Johnson et al. 2002). The number of 

quantitative fire scar histories varies widely among 

ecoregions (Table 3). Both a lack of scientific inquiry 

and the scarce availability of datable fire scarred materials 

contribute to this disparity. Northern Hardwood studies 

are located in Minnesota (Frissell 1993; Clark 1990; 

Heinselman 1973; Spurr 1954), Michigan (Torretti 

2003), Ontario (Dey and Guyette 2000; Cwynar 1977), 

and Vermont (Engstrom and Mann 1991). Central 

Hardwood studies are located in Missouri (Guyette 

and Kabrick 2003; Guyette et al. 2002; Batek et al. 

1999; Guyette and Cutter 1997), Indiana (Guyette et 

al. 2003), Tennessee (Guyette and Stambaugh 2005; 

Harmon 1982; Armbrister 2002), Ohio (Sutherland 

1997), West Virginia (Schuler and McClain 2003), 

Virginia (Sutherland et al. 1992), and Maryland 

(Shumway et al. 2001). Although there are studies for 

the southern pine-oak region (DeVivo 1991, Van Lear 

and Waldrop1989), few are fire history studies from 

fire scars. There are three fire scar sites on the edge 

of the southern pine-oak region in the Lower Boston 

Mountains of Arkansas (Guyette and Spetich 2003). 

Forest-prairie transition sites are on the edge of the Great 

Plains (Guyette and Stambaugh 2005), the oak savannas 

of Wisconsin (Wolf 2004), the oak woodlands and 

savannas of south-central Missouri (Cutter and Guyette 

1994; Guyette and Cutter 1991; Dey et al. 2004), and 

redcedar glades of southwestern Missouri (Guyette and 

McGinnes 1982).

The Warm Continental-Northern Hardwoods region 

has several robust fire histories, though most are in 

forest types with moderate to low oak abundance. 

Several of these were not based on fire scars or derived 

from precisely crossdated tree rings (Whitney 1986; 

Heinselman 1973). Nine studies in this region had 

an average mean fire interval (MFI) of 23 years, with 

a range from 11 to 33 or more years before European 

settlement (ca. 1650 to 1850).

The Hot Continental-Central Hardwoods region is fairly 

well represented, especially in its western half. Here, 

many studies indicate a high frequency of burning in 

oak and oak-pine forests. Fifteen studies had an average 

MFI of 8 years, with a range from 3 to 21or more years 

before European settlement. However, in some areas 

of the Ozarks, high variability in fire frequency can be 

found within small extents (e.g., 3 km) (Guyette et al. 

2002).

Unfortunately, little quantitative fire scar data are 

available for the Subtropical-Southern Pine-Hardwoods 

(Table 3). For this review we used sites (Guyette and 

Spetich 2003) on the edge of this ecoregion in Arkansas. 

Three upland forest sites in Arkansas had an average 

MFI of 11 years, with a range from 5 to 16 or more 

years before European settlement. The topographic 

roughness and low population density of these sites 

during this period probably do not represent the region 

as a whole. However, the southern pine-oak region has 

a number of species and ecosystems with potential fire 

scar data, and represents excellent future opportunities 

for documenting fire history. “Yellow pine” species often 

have resinous wood that preserves well for at least a 

Table 3.—Number of fire scar history studies and sites by Ecoregion

Ecoregion Forest type Studies Sites

Warm continental Northern hardwoods 13 17

Hot continental Central hardwoods 25 60

Subtropical Southern pine hardwoods 1 1

Prairie Forest prairie transition 6 6
2	 Fire 4 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

century after the death of the tree. Live and recently dead 

oaks of the white oak group also can provide excellent 

fire histories.

The forest-prairie transition region is represented by 

several studies in Missouri and Wisconsin, that focus on 

conditions from oak woodlands and savannas to cedar 

glades. Five sites in the forest prairie transition had an 

average MFI of 5 years, with a range from 3 to 8 years or 

more years before European settlement.

The fire scar record (Fig. 1a-c) as indicated by the studies 

cited, provides the timing and magnitude of fire events 

that can be linked to climate. However, the timing of 

annual fire dates is less than exact within a given year 

due to the nature of tree growth and fire scars. Fires 

Figure 1.—Time series of percent trees scarred at sites in or near forests with an oak 

component in Eastern North America. The “complete period site” chronology (a, doted line) 

is compiled from fire scar chronologies that cover the whole or nearly the whole time 

period. The “all sites” chronology (a, solid line) is compiled from all fire scar chronologies 

regardless of length. The eastern chronology (b) is from sites in Ontario, Maryland, Indiana, 

Tennessee, West Virginia, Virginia, and Ohio. The western chronology (c) is from Arkansas, 

Missouri, Wisconsin, Minnesota, and Michigan. See Table 1 for Eastern North American 

site references.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 25

can be dated to the early, mid, and late parts of the 

growing season or to the dormant season (generally 

October through April). Fire scar evidence from eastern 

forests indicates that fires in oak ecosystems occur most 

frequently during the dormant season of tree growth. Of 

the 60 or more study sites in eastern North America in 

which the authors have been involved, about 95 percent 

of the fire scars occurred between annual rings during 

the dormant season. This probably results from surface 

fuel characteristics such as the quantity of leaf litter, its 

density and arrangement, and its potential moisture 

content all of which increased during leafoff and can lead 

to more pyrogenic conditions. The season of burning is 

greatly restricted by primary fine fuels as the controller 

of fire propagation. Surface fuels in deciduous forests 

have more and potentially drier litter when the leaves 

have fallen. Full summer canopies mitigate the loss of 

litter moisture and fuel temperature by decreasing solar 

exposure.

Factors influencing fire intervals

Spatial and Temporal Climate Differences

Climate is a pervasive factor in all fire regimes and has 

complex effects at multiple spatial and temporal scales. 

Spatially, we address regional climate differences as 

well as the effects of year-to-year climate variability on 

fire history sites. The duration of fuel conditioning, 

the duration of snow cover, the length and intensity 

of the warm season, and primary productivity are tied 

to temperature and precipitation. The correlation 

between annual mean maximum temperature (proxy 

period: 1971 to 2000) and pre-European settlement 

mean fire intervals at 38 sites (Table 1) is significant 

(p < 0.001) and strong (r = -0.73, Fig. 2), indicating that 

temperature likely influences many biotic and abiotic 

factors that control fire regimes. Precipitation also was 

significantly correlated (r = -0.56, p < 0.01) with fire 

intervals. The negative correlation value reflects the 

association of shorter intervals between fires occurring at 

sites with higher amounts of precipitation. In some ways 

this is counterintuitive since wetter conditions can create 

shorter fire windows and higher fuel moisture; however, 

increased precipitation generally leads to higher primary 

productivity and potential more rapid fuel accumulation. 

Thus, regional climate parameters, particularly mean 

maximum temperature and precipitation, may have a 

major influence on fire regimes.

Temporally, the effects of yearly to monthly differences 

in climate, particularly drought, are obvious and 

important to fire occurrence and fire regimes. Equally 

important in eastern deciduous forest ecosystems are 

Figure 2.—Mean fire intervals 

(MFI) predicted from proxy mean 

maximum temperature (maxt). 

Mean fire intervals are for the preEuro American period (ca.1650 

to 1850). Temperature is the 

average annual mean maximum 

temperature for 1971 to 2000 minus 

0.4 oC for North American warming. 

Eastern North American sites are 

denoted by “e:” while western North 

American sites are denoted by “w”; 

fire scar data are from 79 sites with 

more than 1,500 trees and 10,000 

fire scars. See Table 1 for Eastern 

North American (e) site references 

and the Model Development 

section of this paper for western 

North American (w) site references. 

The equation in the upper right 

describes the regression line.
2	 Fire 6 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

conditions during leafoff, particularly spring and fall 

season drought. Our preliminary correlation analysis 

of drought and fire extent and severity (percent trees 

scarred) shows that annual reconstructed drought indices 

(Cook et al. 1999) are weakly (r < 0.2) but significantly 

(p < 0.05) associated with the percentage of trees scarred 

at various sites in Eastern North America, particularly 

those in the western portion. Fire occurrence and percent 

trees scarred in eastern oak forests were only weakly 

related to reconstructed annual drought over the past 

three centuries. This probably results from limitations 

of drought reconstructions and their resolution, and 

asynchrony between fire occurrence (dormant season) 

and the bases of reconstructed drought (growing-season 

ring width). However, in years when many sites and 

trees were scarred in Eastern North America, there is 

evidence for widespread drought that continues through 

growing and dormant seasons (Fig. 3). Eight of ten of 

the largest fire years occurred during severe droughts. 

We judge by the percentage of trees scarred (years with 

twice the average percent trees scarred) that major fire 

years in Eastern North America occurred about 3.6 

times per century before fire suppression efforts in forests 

with an oak component. These large fire years often 

are associated with historical documentation (Fig. 3) of 

smoke, wildfire, and fire prevention.

Figure 3.—Percentage of trees scarred at 

sites with fire scar chronologies in Eastern 

North American, illustrating the connection 

between the fire scar record and historical 

documentations of fire history. PDSI is the 

reconstructed Palmer drought severity 

index (Cook et al. 1999). Negative values 

denote increasing drought.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 27

Figure 4.—Regression lines indicate the increased strength of the drought-fire 

relationship during periods of high versus low human population density. Fire data 

are from more than 2500 dated fire scars at 27 sites in the Current River region of the 

Ozark Highlands between 1700 and 1890. Fire area is based on the product of the 

percent of trees and the percent of sites burned annually. PDSI is the reconstructed 

Palmer drought severity index (Cook et al. 1999).

The fire scar record illustrates that drought-fire 

associations are not fully realized in the absence of 

numerous ignitions, as many severe droughts result in 

no fire events. Human population density influences 

the relationship between drought and wildfire by 

increasing ignitions. In fire regimes with few lightning 

ignitions, the effects of drought on fire occurrence 

often are unrealized until human ignitions become 

abundant. For example, in the Ozark Highlands, the 

extent of wildfire is moderately correlated (r = -0.44, 

p = 0.004) with drought during periods with abundant 

anthropogenic ignitions (1850 to 1890) but weakly 

correlated (r = -0.15, p = 0.014) during periods with low 

human population density (1700 to 1849) (Fig. 4). This 

supports the hypothesis that humans may have played a 

significant role in past fire regimes.

When interpreting fire scar records, a common 

assumption is that years with a higher percentage of 

trees scarred result solely from the effects of drought 

on fuels and scarring. However, it is possible that many 

past wildfires were the result of climate-conditioned 

fuels and numerous drought-inspired human ignitions. 

While one might expect an increase in fires owing to 

accidental ignitions during drought periods, there is no 

necessary causation between drought and the number of 

intentional fires. This theory is supported by consistent 

correlations between drought and arson documented by 

the modern fire record. For example, fire season drought 

is significantly correlated with the number of arson fires 

in the Eminence Fire Protection District (1970 to 1989) 

in the Missouri Ozarks (r = -0.76). On a larger scale, 

drought and the number of arson fires summarized by 
2	 Fire 8 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

year and state are significantly correlated (Table 4) (U.S. 

Dep. Agric. 1940-1997). Thus, during much of our 

history, and even during the recent period of societal 

pressure and legal penalties for arson, humans are still 

purposefully burning landscapes during droughts.

Topographic Resistance to Fire Propagation

The frequency of fire is a function of two types of 

ignitions: local and neighboring. For example, fires at 

a site 1 to 3 km2

 in area result from ignitions produced 

locally (new ignitions within the site) or from the 

propagation of ignitions into the site by a spreading fire. 

Thus, fire frequency at a site is both a product of local 

ignition and the resistance of the surrounding landscape 

to the propagation of fire. Part of this resistance is a 

function of topographic features such as hills, valleys, 

and bodies of water that disrupt the continuity of 

fuels (Guyette and Stambaugh, in press). For example, 

the highly dissected topography of the Current River 

watershed in Missouri has been shown to be related 

to the frequency of fires during periods of low human 

population density, when anthropogenic ignitions 

were limited in number (Guyette et al. 2002; Guyette 

and Dey 2000). In topographically rough terrain, 

low-intensity surface fires are impeded by steep slopes, 

discontinuous fuels, and variability in fuel moisture 

and fuel loading. In addition, topographic roughness 

likely mitigates the effect of increasing anthropogenic 

ignitions as many more human ignitions are required to 

maintain the same mean fire interval compared to less 

topographically rough regions (Table 5). Thus, at the 

time of Euro-American settlement, short fire intervals 

(1 to 3 years) in topographically rough landscapes occur 

at later dates as human populations increased. Before 

Table 5.—Effects of topographic roughness on mean fire intervals at increasing human 

population densities (topo-resistance refers to the resistance of a landscape to the propagation 

of fire); details on the calculation of topographic roughness indices found in Guyette et al. 

2002; Peak mean fire intervals are for the first decades of most frequent burning

Site Group Number

sites

ToporesistanceRoughness 

index MFI at peak

Population at 

peak no./km2 Year at peak

Boston Mts., AR 6 high 1.08 2.7 5.5 1875

Current River, MO 10 moderate 1.020 3.5 0.64 1850

Highland Rim, TN 4 low 1.0004 2.3 0.50 1770

Table 4.—Relationship between number of arson fires and drought by 

state (Drought data are the annual Palmer Drought Severity Index; the 

lower the value of this index, the more severe the drought; thus, the 

negative correlation between drought and arson; correlation coefficients 

in bold are significant at (p<0.05); fire data: U.S. Dep. Agric. 1940-1997)

State Correlation

(No. arson fires 

and drought)

Mean no. 

arson fires

Mean no.

lightning

fires

No. of 

years of 

data

Missouri -0.39 1069 20 48

Arkansas -0.61 1638 85 48

Illinois -0.49 46 0.72 48

Indiana -0.29 36 1.7 48

Tennessee 0.04 1148 12 48

S. Carolina -0.36 1918 58 48

Florida -0.42 2769 580 48
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 29

1875, about 10 times as many humans were required 

to maintain a fire frequency of less than 3 years in the 

topographically rough Boston Mountains, Arkansas, as 

were required in the Oak Barrens of the Highland Rim 

in middle Tennessee (Table 5).

Fire and Culture

Of all of the cultural aspects affecting wildland fire, 

none have been as profound and effective as suppression 

efforts enabled by the modern technologies of ground 

and air transportation. Although grazing, landscape fuel 

fragmentation, intentional burning, and other cultural 

influences have been detected in fire scar records, these 

effects are not as pronounced as fire suppression in 

modern industrial societies. All but a handful of fire scar 

chronologies show a significant decline in fire frequency 

in the mid to late 20th century. The few fire scar 

chronologies that do not have a decline in fire frequency 

often are derived from private lands with owners who 

have a tradition of burning, or are in remote regions with 

long response times in fire protection (Jenkins 1997).

Before fire suppression, cultural values concerning 

wildland burning are not evident in the fire scar record 

and may be secondary to increasing anthropogenic 

ignitions as a function of human population density. In 

many environments, humans are a fire obligate species. 

Accidental ignitions, purposeful ignitions, machine 

fires, and debris burning are a few of the hundreds of 

human activities that can result in wildland fires. The 

sheer number of humans in an ecosystem may be more 

important than their cultural attitudes about the use 

of fire. Unlike fire frequency, which is tempered by fire 

suppression, fuel fragmentation, and reduction, the 

number of ignitions continues to rise with increasing 

human population.

The fire scar record suggests that Homo sapiens is a 

keystone species in many fire regimes. A keystone 

species influences the “environmental balance of an 

area or habitat” through a chain of events (e.g., trophic, 

reproductive, or abiotic modifications). Keystone 

species have a disproportionate influence on community 

structure in excess of their abundance. The fire scar 

record indicates that even at low population densities, 

humans had a profound effect on fire regimes and 

vegetation (Batek et al. 1999; Guyette and Kabrick 

2003). The influence of a keystone species is derived 

from the ability to control the most competitive species 

(i.e., tree species). Temporally and spatially, fire scar 

frequency has been associated with the presence of 

humans. The absence of a keystone species releases 

highly competitive plants (trees) from the limiting effects 

of fire and allows them to exclude other plants and 

their herbivores. In forested regions, fire scar frequency 

increases rapidly as humans culture the forest for both 

wild and domestic large herbivores. The theory that 

humans are a keystone species in fire regimes may be 

especially relevant in Eastern North America, where 

climate windows suitable for burning often are short and 

the probability of natural ignitions is low. The fire scar 

record in Eastern North America is only weakly related 

to the abundance of lightning fires as documented later 

in this paper. Unlike lightning, humans can target their 

ignitions with respect to fuel conditions and location.

Population Density and Anthropogenic 

Ignition Rates

Since the entry of Homo sapiens into North America 

more than 12,000 years ago, humans have been the 

dominant ignition factor in many ecosystems. Although 

some have argued that there were not enough humans 

to impact flora and fauna at regional levels, the fire scar 

evidence suggests otherwise. Few humans are necessary 

to add a significant level of fire disturbance. The 

topographically rough Missouri Ozarks reach ignition 

saturation at about 0.64 human per km2 or less (Guyette 

and Dey 2000). The rougher Boston Mountains of 

Arkansas reach ignition saturation at about four or five 

humans per km2. The smooth oak flats of the Highland 

Rim in middle Tennessee reach ignition saturation 

at about 0.50 human per km2. Even at relatively low 

human population densities in a moderately dissected 

landscape (e.g., Missouri Ozarks) mean fire intervals of 

10 years were supported by populations of less than 0.10 

human per km2 during a “depopulated” period (1680 to 

1800).

A theoretical analysis of scenarios of human population 

density, ignition type, and culture illustrates the potential 

influence of anthropogenic ignitions in past fire regimes 

and consistency with the fire scar record. The population 
30	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

of North America (north of Mexico) has been estimated 

to be from 1 million to more than 10 million at the 

time of first contact and before population reductions 

by a variety of agents, e.g., introduced diseases. We 

chose the conservative estimate of about 1 million 

humans for this analysis. We used 500,000 humans for 

the estimate of population in the Eastern United States 

and southeastern Canada. The study area in eastern 

North America is about 3,024,000 km2

. We then 

subtracted the area of the Great Lakes (245,000 km2

) 

to obtain an area of 2,779,000 km2

. Although rough, 

this estimate is more precise than population estimates. 

Thus, population density roughly averaged 1 person 

for every 5.5 km2

, or 0.18 human per km2 (500,000 

humans/2,779,000 km2

). Given this population 

density, we generate quantitative scenarios of the rates 

of purposeful and accidental ignitions. Present-day rates 

of human-caused fires in forested areas of the Missouri 

Ozarks average one wildfire per 350 humans per year 

(0.0029 fire/human/yr). This rate occurred during an 

era of fire suppression (1991 to 2003), laws and societal 

pressures against burning, relatively few open fires (e.g., 

camp fires) per person, and limits on fire propagation 

imposed by land fragmentation. This rate applies to 

all arson (31 percent) and accidental fires (66 percent). 

Prior to European settlement, if the people of a culture 

viewed fire neutrally, the rate of ignitions by humans 

might be 10 times this amount (0.029 fires/human/yr). 

Thus, based on a pre-European settlement population 

and a neutral fire culture, the rate for human ignitions 

would be 0.0052 human-caused fires per km2

 per year 

(i.e., 0.029 fire/human/yr multiplied by 0.18 human 

per/km2). This translates to a single human ignition 

for every 190 km2

 per year. If the people of a culture 

purposely used wildland fire as a tool (for many possible 

reasons) the rate of human ignitions might be 100 times 

greater, or 0.052 human ignition per km2

/year (i.e., 

0.29 fire/human/yr multiplied by 0.18 human per/km2

). 

This translates to a single human ignition for every 19 

km2 per year. This rate (0.052 human-caused fire/km2/

yr) is about 200 times the rate of lightning ignition in 

the Central Hardwoods region (0.00025/ km2

/year) 

(Schroeder and Buck 1970). For example, if the average 

fire size was 1.9 km2

, this rate of ignition would maintain 

a fire rotation interval of about 10 years. This scenario 

and fire interval is not unlike the rate of burning (13 

percent sites scarred annually) that occurred in 1790 

in the moderately rough Current River landscape at 

population levels of about 0.017 human/km2

 (Guyette et 

al. 2002). The anthropogenic ignition rate is potentially 

much greater if intentional burning occurs (Williams 

2000, Williams 2001). We conclude that even low levels 

of human population density combined with a culture 

of landscape burning were sufficient to provide levels 

of ignitions needed to maintain the fire frequencies we 

document with fire scars in Eastern North America.

Synthesis and modeling of

Pre-European fire intervals

Quantitative and empirically derived models of fire 

regimes can be used in many ways. Models allow past 

fire regimes to be estimated for ecosystems that have 

no on-site fire history information. Models can be used 

in conjunction with soil, geology, and species data to 

reconstruct past and potential flora and fauna. Thus, 

land managers interested in returning ecosystems to preEuropean settlement conditions (including fuels) using 

prescribed fire can utilize models to estimate components 

of fire regimes including fire frequency. Researchers 

can use equations of this type to create a continuous 

landscape overlay of past fire frequency. Models without 

vegetation input also can be useful for independently 

estimating fire frequency at research vegetation plots, and 

for making inferences about possible changes in future 

fire regimes due to changes in population and climate. 

The preliminary results of the modeling that follows 

should be used with caution as they are temporally and 

spatially coarse scale, requiring additional validation 

and data from many regions of Eastern North America. 

Perhaps the most important aspect of fire interval 

models and fire history may not be the information 

provided on fire regimes but the perspective provided on 

the long-term interactions between humans and their 

environment.

Model Development

A preliminary model that predicts mean fire intervals for 

eastern oak forest ecosystems before European settlement 

was developed from fire scar data. The first iteration of 

the model was empirically derived from available fire 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 31

scar data at 38 sites in Eastern North America (Table 

1, col. 5). Although this data set showed predictive 

power, the eastern fire history data alone was inadequate 

in the range and distribution of the variables and did 

not meet the assumption of statistical normality. Thus, 

we used pre-European fire scar interval data from an 

additional 41 sites in the Western United States (Brown 

and Sieg 1996; Brown et al. 1997; Caprio 1998; Caprio 

and Swetnam 1995; Donnegan et al. 2001; Finny and 

Martin 1989; Fulé et al. 2003; Heyerdahl et al. 2001; 

Kipfmueller 2003; Miller and Rose 1999; Moir 1982; 

Swetnam et al. 1989 ; 1991).

These additional data extended the range of temperature 

and fire interval observations, increased the normality 

of the distribution of temperature means, more than 

doubled the degrees of freedom, and served to examine 

the hypothesis and model under a much broader set 

of conditions. We used published mean fire intervals, 

though in several cases we divided the period of record 

by the number of fires to estimate mean fire intervals. 

This was necessary when data were not expressed as 

mean fire intervals, data were not presented in FHX2

format (Grissino-Mayer 2001), fire dates were not 

published, long fire intervals were open ended, or the 

fire scar dates were not crossdated. Since the variables we 

used in this analysis are being refined and additional fire 

data are needed and forthcoming in many regions, we 

present this model as a work in progress with updated 

versions expected in the future. The two predictor 

variables of mean fire intervals are mean maximum 

temperature (maxt) and human population density 

(pop). Multicollinearity among predictor variables was 

negligible. Of the possible intercorrelations among 

variables, none was significant. The largest variance 

inflation factor (VIF) was 1.19, well below a VIF 

of 10, which would indicate that multicollinearity 

was influencing least squares estimates. The model is 

described by the regression equation:

MFI = -12.7 + 64.5e-0.098*maxt - (6.3*ln [pop]),

 (Equation 1)

where MFI is the mean fire interval, maxt is the proxy 

mean maximum temperature (30 year average in 

degrees Celsius), 

ln[pop] is the natural log of human population 

density (humans per km2

), 

df = 78, r2

 = 0.75, all variables are significant 

(p <0.01).

We tested the model’s stability and predictive power by 

sampling half data with replacement 30 times. Variance 

explained (r2) ranged from 0.63 to 0.83, the standard 

deviation of the coefficient of determination was 

0.05; the r2 mode was 0.75. The model and predictor 

variables tested significant (p < 0.01) in all model runs. 

The model predictions did not rely on extreme or rare 

sets of observations and did not vary greatly by the set 

of observations tested. On the basis of these results, 

the predictive power of the model as measured by the 

coefficient of determination proved stable.

The model predicted the mean fire intervals (Fig. 5) 

for the presence of fire in a 1- to 3-km2

 area during the 

150 years before Euro-American settlement. Due to 

a westward progression of European settlement, this 

period varies by settlement date and the length of the 

fire scar record of each site, but generally begins from 

about 1650 to 1750 and ends between 1780 and 1850. 

The prediction period for this model also was restricted 

because the relationship between humans and fire scar 

frequency was positive only when ignitions were limited 

(Guyette et al. 2002). During later periods when human 

population density is high, this relationship becomes 

more complex and may even be negative. Also, the 

regression model does not predict or include stand 

replacement fires (the data are based entirely on survivor 

trees) or extend to prairies or regions where fire intervals 

are longer than the lives of recorder trees. 

The useful components (predictor variables) of this 

preliminary model and map also are its caveats. One 

of the advantages of this model is that it is based on 

variables that change and can be used to examine 

different scenarios, for example, landscapes without 

humans or possible effects of climatic warming or 

cooling. One disadvantage is that spatial predictions 

probably are more static than actual (and often 

unknown) conditions due to the difficulty of variable 

quantification. This is especially true for the effects 
3	 Fire 2 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

of human-caused ignitions, which can be highly and 

abruptly variable through time and space.

Mean Maximum Temperature Proxy

We used annual mean maximum temperature (Daly et 

al. 2004), a precisely measured and modeled quantity 

compared to fire intervals, as a predictor of mean fire 

interval in Eastern North America. Because our period of 

interest (pre-Euro American settlement) for fire history 

and mean maximum temperature (1971 to 2000) are 

not temporally matched, we subtracted 0.4oC from the 

mean maximum temperature values to compensate for 

the effects of recent warming (Mann et al. 1998). Error 

caused by the non-temporal overlap of the temperature 

record with fire scar history will be minimal because 

the scale of temperature change between 1750 and 

1985 (0.4o

C) is small (about 35 times less) compared to 

differences among sites (about 14o

C). Mean maximum 

temperature probably influences and controls many 

biotic and abiotic components that influence fire regimes 

and fire intervals such as fuel moisture, fuel production, 

and combustion. For example, the length of the fire 

season, as determined by the number of months in 

which 90 percent of the acres are burned in the four oak 

Figure 5.—First approximation of coarse scale mean fire intervals in the Eastern United States as 

predicted by mean maximum temperature and human population density (Equation 1). The mapped 

mean fire intervals are based on fire scar histories and model results generated from forested 

landscapes and nonstand replacement fire events. Mapped estimates are not valid for wetlands, 

grasslands, and other vegetation types where trees do not grow. The pre-Euro American period is 

based on fire scar intervals between 1650 and 1850. Estimates near ocean edges are unclassified 

(black). Mapped mean fire interval estimates are only as stable as human populations (e.g., several 

centuries earlier, the concentrated populations of the Mississippian cultural phase would have 

shortened fire intervals in the Mississippi and Ohio River Valleys).
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 33

ecoregions (National Fire Occurrence Database and GIS 

Coverage 2005), is strongly correlated with maximum 

temperature (r = 0.77, p = 0.01). We speculate that 

mean maximum temperature is related to fire intervals 

and regimes in many ways, including:

1. The length of the fire season as influenced 

by snow cover at northern latitudes and highelevation sites.

2. The length of the fire season as determined by 

the drying rate of fuels.

3. The direct effect of temperature in combustion 

reactions.

4. Mean maximum temperatures most often occur 

during the most fire-prone time of day. 

5. The amount, types, and decay rates of fuels in 

an ecosystem are determined in part by 

temperature.

Other basic climate variables were considered as 

predictors. Precipitation, mean minimum temperature, 

and mean average temperature were included in 

the analysis but did not enter the stepwise multiple 

regression equation as significant. The fact that 

precipitation did not enter the model in stepwise 

regression probably reflects the potential complexity 

of the response of fire regimes to precipitation in hot 

versus cool climates. For instance, fire can be frequent in 

both hot-wet (Florida) and hot-dry (southern Arizona) 

climates. Also, we found that lightning ignition rates 

based on generalized maps of lighting ignitions of forest 

fires (Schroeder and Buck 1970) did not enter the 

stepwise regression, and that correlations between mean 

fire intervals and lightning ignitions were not significant 

(r = 0.16, p = 0.16). However, separation of the data 

into eastern and western sites yielded a significant 

correlation (r = 0.40, p < 0.01) with lightning fires in 

Western but not Eastern North America (r = 0.12, p = 

0.46). This result partially reflects the lack of fire history 

sites in the in the lightning and fire-prone Southeastern 

United States.

Population Density and Ignition Proxy

We used human population density as a proxy for 

the number of anthropogenic ignitions. Although the 

estimation of Native American populations is fraught 

with problems (Henige 1998), progress has been made 

in assessing estimates and observations made by early 

workers (Denevan 1992; Mooney1928). We used 

mapped population density classes of Native Americans 

in North America (Driver and Massey 1957) and more 

detailed population estimates in the Ozarks (Guyette 

et al. 2002). We used population class variables to 

facilitate the quantification of pre-census estimates of 

human population density, which have a large degree of 

potential error and may never be known precisely. These 

estimates are consistent with conservative estimates 

of Native American population (Waldman 1985). 

However, temporal continuity in population density 

estimates is variable. Driver and Massey’s mapping 

of human population is not unlike the relative spatial 

distribution of human population density in 2000 (U.S. 

Census Bur. 2000), illustrating that suitable human 

habitat (and spatially relative population densities) 

is somewhat consistent through time. For example, 

population densities are high during both periods along 

coastal areas, in the Northeast, near the Great Lakes, in 

localized areas of the Southwest, and near Appalachia. 

Also, spatial error in population estimates may be no 

greater than the variance in population density owing to 

the migration, immigration, and decline of populations 

of humans (Thornton 1987) during the two centuries 

of the pre-Euro American settlement fire scar record. 

Although these factors make precise calibration between 

fire frequency and population density difficult over large 

temporal and spatial scales, their significant (p < 0.01) 

correlation (r = - 0.45) supports the theory that humans 

had large-scale effects on fire regimes.

Mapping Fire Intervals

The model results were mapped using ESRI®ArcGISTM

software (Environ. Syst. Res. Inst. 2005). Gridded 

mean maximum temperature data (Daly et al. 2004) 

and coverage of human population density (Driver 

and Massey 1957) were applied to Equation 1 to 

produce estimates of mean fire intervals for a pre-Euro 

American settlement period (about 1650 to 1850). 
34	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Prior to mapping, population data were smoothed 

using a circular (25-km radius) neighborhood mean to 

more closely reflect the mobility of humans. Mapped 

coarse scale fire intervals have error based on the model 

(Equation 1). The model error indicates that the 95-

percent model confidence interval is about ±3 years while 

the 95-percent prediction interval is about ±11 years 

(less for shorter intervals). These error estimates include 

all the variability in vegetation, topography, and local 

human population not evident in the predictor variables 

that can affect fire frequency at finer spatial scales. 

During periods that precede model calibration, the 

model may work within limits but the mapped estimates 

of mean fire intervals must be based on temporally 

appropriate predictor variable data.

Conclusion

The quantitative history of wildland fire derived from 

fire scar studies in Eastern North America provides 

several insights into past and future fire regimes. 

Seemingly important variables such as precipitation did 

not explain variance in mean fire intervals beyond what 

was explained by temperature. Variables relevant to fire 

regimes and fire intervals in eastern oak forests were 

identified and include mean maximum temperature, 

human population density, extreme drought events, and 

topographic resistance to the spread of fire. Some of 

these variables are dynamic, such as human population 

and temperature, and their potential change in the future 

may influence fire regimes. Understanding how these 

variables interact to influence fire regimes will aid in 

assessing future fire risk with changes in fuel continuity, 

temperature, extreme climate events, human culture, and 

population density.

The interaction between drought and intentional human 

ignitions will make future fire regimes potentially 

unstable and difficult to predict. Human populations 

are increasing and will provide more ignitions, making 

fire regimes more responsive to drought. This will be 

countered to some extent by improved suppression 

response time and new technologies. The stability of fire 

regimes in changing climates may be low, particularly 

in landscapes with a low resistance to fire propagation 

and changes in the number and distribution of human 

ignitions. Drought has inspired purposeful human 

burning both in the past and present. It is doubtful that 

this relationship will cease to exist in the future.

The effects of topography on the propagation of fire 

have been understood by humans for a long time. Fire 

scar data quantified this effect at a general landscape 

level. The mitigation of fire by topography has been 

relatively static over geologic time scales compared to the 

ephemeral changes caused by the frequency of human 

ignitions. Thus, in terms of temporal scale, topographic 

roughness is by far one of the most consistent and 

important variables affecting the frequency of fire.

Fire history modeling and mapping from fire scars 

can provide estimates of fire intervals for restoration 

and reference conditions (Maclean and Cleland 2003; 

McKenzie et al. 2000; Morgan et al. 2001). However, 

the quality of any model or map is dependent on the 

quality of the data from which it is derived. In this study, 

we mapped large regions of the Southern and Eastern 

United States for which pre-European settlement fire scar 

histories were unavailable. Therefore, estimates for these 

regions are uncalibrated and approximate but might be 

improved by future studies and data collection.

Acknowledgments

The authors thank all of the investigators whose fire 

histories we used in this article, in particular, the 

excellent Michigan fire history work by Rebecca Torretti 

and advisor Alan Rebertus. This review and analysis 

was made possible by support from the USDA Forest 

Service’s North Central and Southern Research Stations, 

National Park Service, Ontario Ministry of Natural 

Resources, Missouri Department of Conservation, 

CH2MHILL, and the U.S. Air Force.

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40	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Understanding the Evidence for Historical Fire Across

Eastern Forests

Charles M. Ruffner1

Abstract.—Evidence for historical fire across the eastern deciduous biome spans several fields, including 

paleoecology, fire scar analysis, witness tree studies, historical documents and ethnographic sources. 

In this paper I provide an overview of many of these methods as well as the limitations and examples 

of each. While the use of any single approach has its cautions and pitfalls, the interpretation of the 

considerable material available allows a view into the history and effects of fire on eastern oak forests.

conditions for fire so that land managers can begin to 

assemble and interpret evidence for their respective 

holdings, no matter where they occur across the eastern 

forests. I hope that many engaged land managers will 

embrace the historical ecology approach in researching 

and managing their properties. Because so many 

excellent reviews have been published I reference them 

often so that attendees here can find those key resources 

and use the many citations included.

Investigating the role of fire in eastern 

oak forests

To a large extent, historical fire has been implicated 

in the wide distribution of oak forest and woodlands, 

prairies, savannas, and glades that are found within 

nearly every state in the eastern deciduous biome (see 

reviews by Lorimer 1985; Abrams 1992; 2002, Van Lear 

and Watt 1993; Whitney 1994; Dey 2002). The wide 

range of evidence concerning historical fire across many 

parts of the East ranges from paleoecology studies (see 

Patterson, this volume), fire histories and chronologies 

(Guyette, this volume), archaeological investigations 

(Chapman et al. 1982; Delcourt et al. 1986, 1998), 

native American sources (Parker 1968’ Adovasio et 

al. 1985’ Doolittle 1992; Kimmerer and Lake 2001), 

and witness tree analysis and surveyor records (Bourdo 

1956; Hutchison 1988; Whitney 1994; Black and 

Abrams 2001; Abrams 2003), to historical documents 

such as deed descriptions and tax records (Foster 1992; 

Olson 1996), plat maps, travel documents and journals 

(Schoolcraft 1820; Burges 1965; Ladd 1991,;Whitney 

1994; Olson 1996; Ruffner and Arabas 1999), and stand 

structure analysis and dendroecological studies (Henry 

and Swan 1977; Dorney and Dorney 1989; Nowacki 

and Abrams 1992; Ruffner and Abrams 2002).

Introduction

As foresters, ecologists, and land managers, each of us 

is charged with understanding our ecosystems and the 

processes that have acted to form those systems through 

time. Learning as much as possible about the original 

vegetation and structure of our systems is vital to 

identify and manage these natural areas for the optimal 

biodiversity inherent in original conditions (Loucks 

1970; Ladd 1991; Kay 1995; Olson 1996). Across the 

East, the historical role of fire has been implicated in 

driving diversity and maintaining keystone species in 

many ecosystems and the body of literature investigating 

these processes continues to grow. Mutch (1991) 

suggested that fire regime information is “absolutely 

essential background data for the appropriate design 

and implementation of resource management projects 

at the ecosystem and landscape level of organization.” 

In southern Illinois, I work with landowners and 

agency managers in finding information concerning 

the pre- and post-European settlement conditions of 

their land holdings. These endeavors often focus on 

gaining a better understanding of the historic range of 

variability represented in some systems, or understanding 

the spatial distribution of hill prairies and barrens. No 

matter the objective, much of the data sources available 

for these studies are found across several disciplines and 

thus are sometimes difficult to access or, quite frankly, 

understand.

The objectives of this paper are to review many of the 

basic approaches used to interpret or infer historical 

1

Department of Forestry, Southern Illinois University, 

Carbondale, IL 62901-4411, 618-453-7469, email: 

Ruffner@siu.edu
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 41

Specific limitations and/or assumptions about many 

of these data formats must be considered (Forman 

and Russell 1983, also see review in Whitney 1994). 

For many of the primary documents, the historical 

context during its writing or surveyor bias against some 

landscape feature or misidentification of a species and 

least of all the observers’ training and skill level may 

shade the interpretation of these historic documents. 

Nonetheless, as many authors note, these data sources 

often are the only representation of the pre-European 

landscape. Despite their variable training levels, many 

primary authors had “ the unsurpassable advantage 

of actually having seen presettlement landscapes and 

aboriginal land management practices” (Ladd 1991). 

Thus, while many methods have their critics, when 

interpreted together or by applying an historical 

ecological approach they provide the land manager with 

much needed information to place her/his land parcel 

in its historical context and begin to model the types of 

vegetation structure and function appropriate for that 

area (Myers and Peroni 1983).

Paleoecology

Across the eastern deciduous biome, many authors have 

used pollen and charcoal analysis to reconstruct and 

interpret vegetational development at different scales 

(Watts 1979; Webb 1981; Whitney 1994). According 

to Patterson (this volume) some areas of the region have 

fairly well documented paleoecological reconstructions 

based on the work of key researchers, for example, Watts’ 

work in the Appalachians (Watts 1979), Davis’ work 

in the northeast and Lake States (Davis 1983), Clark’s 

work in the Midwest and Northeast, and work in New 

England, primarily from Patterson and associates at the 

University of Massachusetts and from Foster and others 

at Harvard Forest.

In comparing sedimentary charcoal and archaeological 

site distributions in New England, Patterson and 

Sassaman (1988) found that fires were common on 

coastal sites where native populations were greatest 

and their land-use practices most intensive. Across the 

region, others have noted archaeological site distributions 

corresponding well with areas characterized by high 

oak pollen percentages and usually elevated amounts of 

charcoal fragments (Dincauze and Mulholland 1977; 

Patterson and Sassaman 1988; Delcourt et al. 1998; 

Fuller et al. 1998). More recently, Clark and Royall 

(1995) contributed greatly to this discussion reporting 

a transition from northern hardwood to white pineoak forests at Crawford Lake, Ontario, following a 

period of Iroquois occupation and agricultural clearing. 

Moreover, Ceci (1979) and Loeb (1998) discussed native 

corn and hickory planting episodes from the pollen 

record in the coastal regions of New York. Delcourt et 

al. (1998) reported that Archaic and Woodland period 

natives cleared forest gaps to cultivate native plants, 

and that human-caused fires increased populations of 

fire tolerant oaks, chestnut, and pine in upland forests 

of the Cumberland Plateau of central Kentucky. Still, 

Clark and others have provided evidence that some oak 

communities can be maintained in oak-chestnut with 

litle charcoal present in their sedimentary records (Clark 

and Royall 1995; Foster et al. 2002). While several 

regional studies suggest the correlation of fire occurrence 

and native occupation with oak forest distribution, 

more research at the local level must be completed to 

better understand the pre-European landscape across the 

Northeast (Black and Abrams 2001; Parshall and Foster 

2002).

Fire Scars

Another quantitative approach to understanding 

historical fire is using dating fire scars or tree-ring 

analysis to reconstruct disturbance regimes at native 

sites (Batek et al. 1999; Shumway et al. 2001; Ruffner 

and Abrams 2002). Few presettlement fire papers 

have been published for the Northeast; most represent 

fire evidence from the Midwest and Central plains 

(Abrams 1985; Guyette and Cutter 1991; Cutter and 

Guyette1994; Batek et al. 1999). Buell et al. (1954) 

reported six fires scars from a single tree dating from 

1641 to 1711 in Mettler’s Woods, New Jersey, indicated 

a mean fire interval (time between fires) of 14 years. 

In western Maryland, Shumway et al. (2001) reported 

a mean fire interval (for consistency) of 7.6 years in 

an old-growth upland oak-hickory forest. Across the 

region, estimates of mean fire interval within oak forests 

are surprisingly similar ranging from 2 to 24 years 

(Shumway et al. 2001). At many sites, fire frequency 

increased or decreased during the initial period following 

European settlement depending on the stand and 
4	 Fire 2 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

region. Fire histories for the Missouri Ozarks indicate 

that during periods of late Native American settlement 

(1701 to 1820), fire return intervals were longer (11.96 

years ± 2.4, mean ± SE) than those during European 

settlement (3.64 years ± 0.35) (adapted from Guyette 

and Cutter 1991). In southern Illinois, little is known 

of the historical fire regime because most of the needed 

fire scars were lost after logging of the primary forests 

and deterioration of cut stumps (Robertson and Heikens 

1994; Olson 1996). Robertson and Heikens (1994) 

reported high fire ignitions during European settlement 

due to farmers clearing underbrush from the forest, 

closely following Guyette and Dey’s model of frequent 

European ignitions immediately following settlement.

Moving eastward, a fire history study of mixed 

oak forests originating after 1850 in southeastern 

Ohio revealed that fire return interval averaged 

7.5 years (Sutherland 1997). In northwestern 

Pennsylvania, Ruffner and Abrams (2002) reported 

dendrochronological evidence at a native Seneca village 

site revealed a frequent, low-intensity fire regime with a 

return interval of 11 to 26 years during native habitation. 

However, following European settlement (post-1800) the 

fire free period is significantly longer with a disturbancefree interval of 28.5 ± 2.8 years. Where prehistoric fire 

scars are available, these studies have provided valuable 

insight into native disturbances and their localized 

impacts. However, they are largely restricted to the area 

immediately surrounding the study sites and cannot 

characterize native influences at the landscape level 

(Batek et al. 1999).

Witness Trees

During the period of early European settlement in the 

Eastern United States, land surveyors recorded and 

marked witness trees (warrant, bearing, and/or corner) 

to identify property corners and boundaries (Lutz 

1930; Spurr 1951; Bourdo 1956). When compiled 

from surveyors’ notes, these trees provide a landscape 

level snapshot of forest composition at the time of 

European settlement and have been used to reconstruct 

regional vegetation composition throughout the Eastern 

and Midwestern United States (Lutz 1930; Siccama 

1971; Whitney 1986; 1990, Loeb 1987; Seischab 

1990, 1992; Marks and Gardescu 1992; Nowacki 

and Abrams 1992; Abrams and Ruffner 1995; Black 

and Abrams 2001). Such vegetation reconstructions 

are valuable in areas where most original vegetation 

has been destroyed or altered, making it difficult to 

assess natural vegetation types or disturbance regimes. 

In many studies, reconstructed patterns of vegetation 

have been attributed to edaphic conditions such as 

topography, soils, slope class or drainage classes (Gordon 

1940; Abrams and Ruffner 1995). In more recent years 

studies have taken a more integrative approach and 

recognized that native populations also may have affected 

vegetation patterns. A study by Dorney and Dorney 

(1989) documented a native maintained oak savanna 

in northeastern Wisconsin using General Land Office 

survey information.

In documenting pre-European vegetation of central New 

York State, Marks and Gardescu (1992) identified old 

Native American clearings and reported oak-hickorypine growing in areas previously inhabited by peoples of 

the Iroquois Confederacy. In western New York, on lands 

surveyed for the Holland Land Company, including 

Gordon’s (1940) area, Seischab (1990) reported 

that most oak communities occurred in southern 

areas along the Allegheny River and its tributaries, 

dominating a landscape formerly inhabited by Seneca 

people of the Iroquois Confederacy. Within the Lower 

Susquehanna Valley of Pennsylvania, Black and Abrams 

(2001) reported significant differences in witness tree 

distributions between areas adjacent to Native American 

village sites and those in areas with low native activity.

Whitney and Decant (2003) suggested that Iroquois 

habitation may be associated with oak-chestnut forests 

along the French Creek area in their witness tree study of 

northwestern Pennsylvania.

Most recently, Black et al. (unpublished) integrated 

witness tree distributions, Native American archaeology, 

and geologic and topographic variables to investigate the 

relationships between Native American populations and 

pre-European settlement forest types on the Allegheny 

Plateau of northwestern Pennsylvania. Logistic regression 

of natural and cultural variables demonstrated that 

Native American habitation and travelways were the 

most significant predictor of oak, hickory, and chestnut 

trees in the pre-settlement forest. Although cause and 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 43

effect cannot be tested at this time (the paleoecology 

data are still being analyzed), the authors suggest that 

long-term Native American activities such as agricultural 

clearing and burning, as well as possible wood and mast 

resource extraction selected for the disturbance, adapted 

oak-hickory and chestnut. Overall, an increasing number 

of studies suggest that Native Americans had at least 

some association with forest composition, though the 

degree of the relationship often is not quantified.

Historical documents

Much of the literature surrounding Native American 

effects on vegetation have been developed from some 

type of historical document. By the middle of the 20th 

century, several key historical geographers had laid 

the framework for investigating native land uses e.g., 

Maxwell (1910), Bromley (1935), Day (1953), Stewart 

(1963), and Sauer (1975). These authors reviewed 

numerous primary and secondary sources to build their 

cases that native groups had influenced vegetation to 

varying degrees across the continent (Whitney 1994). 

While some authors suggested the widespread use 

of fire by natives, others cautioned against the idea 

of natives wantonly burning the landscape. Still, the 

preponderance of early forests described as “open and 

park like” led Bromley (1935) to conclude that it was 

“due to the universal factor of fire, fostered by the 

original inhabitants to facilitate travel and hunting” 

(Nowacki 2002). Indeed, natives affected ecosystem 

development in a variety of ways: hunting, fishing, 

agricultural clearing, and associated activities (including 

the introduction and dissemination of new cultigens), 

wood gathering, village and trail construction, and 

habitat manipulation (Maxwell 1910; Day 1953; Mellars 

1976; Nowacki 2002). It is widely accepted that natives 

intentionally used fire for more than 70 reasons (Lewis 

1993), particularly in the alteration and maintenance of 

surrounding environments for their benefit (Day 1953; 

Cronon 1983; Mellars 1976; Doolittle 2004).

Examples from early travel documents include William 

Strachey’s (1620) description of the Hampton, Virginia, 

area as “the seat sometime of a thowsand Indians and 

three hundred houses...which is the reason that so much 

ground is there cliered and opened, enough already 

prepared to receive corne and make viniards of two or 

three thowsand acres.” Another published in 1672, The 

Discoveries of John Lederer: In three several marches 

from Virginia to the west of Carolina, relates open 

valleys and active burning of pine forests near Indian 

settlements throughout the Piedmont and Ridge and 

Valley provinces. In 1794, William Bartram traveled 

through the Cherokee, Creek, and Choctaw nations, 

characterizing their farms, orchards, and woodlands. In 

1810, Francois Michaux published memoirs of his and 

his father’s travels through the southern Appalachians 

with their astute depictions of native village life and 

culture. Farther west, Henry Schoolcraft’s 1821 Ozark 

Journals documents his travels through the Ozark 

region where his training allowed him to accurately 

identify plant assemblages and report on the character 

and conditions of many unique ecological sites and 

phenomenon encountered on his trips through Indian 

territory. Most authors recorded an anthropogenic 

landscape heavily influenced by native activities and 

populations.

Researchers using surveyor accounts, treaties, deeds, 

wills, sheriff sales, tax records, and other collected papers 

and manuscripts have characterized and explained 

vegetation change from pre-European settlement to 

the present (Raup 1966; Cronon 1983; Foster 1992; 

Orwig and Abrams 1994; Ruffner and Abrams 1998; 

Foster 2002). While native effects differ across these 

studies, there are general patterns. For instance, many 

early documents reflect a managed landscape with 

tended agricultural fields interspersed through open 

woodlands and numerous abandoned habitation areas. 

This clearly suggests a cultural landscape. In a search 

of historical documents of Cape Cod, one of the 

earliest places written about by Europeans, Patterson 

and Ruffner found evidence for large “open fields” 

and “upland meadows” surrounding native settlements 

(Plymouth Colony records, vol. 3, p. 85). A 1648 deed 

records “the fields over the pond have been cleared 

and improved by the Indians as long as the oldest can 

remember” (Massachusetts Archives. 33: 245-247). As 

in other places across the East, these “planting fields” 

were the most commonly sought after real estate by 

Freemen during the ensuing years of European expansion 

(Williams 1989, Whitney 1994). 
44	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Another common observation is the usefulness and 

purposeful nature of many native burning regimes, 

which usually were adopted by incoming European 

settlers (Whitney 1994). Whether the reason was 

reducing rattlesnake and insect populations, improving 

browse or fruit production, or easing travel through the 

forest, authors have reported the widespread adoption of 

the native practices. Plymouth Colony records revealed 

evidence for early prescribed burning of pine lands by 

selected officials because they recognized the need to 

reduce potential fire hazards around settlements and 

improvements (Lovell 1984).

Conclusion

The use of historical evidence to reconstruct past 

vegetation distribution and fire regimes spans 

paleoecological, ethnobotanical, and archaeological 

fields. While the field of historical ecology has grown 

over the last 50 years to include many disciplines and 

journal outlets, I hope that this introduction contains 

sufficient examples of outstanding research and review 

articles to allow newcomers to find new sources of 

information. It is testament to the usefulness of these 

historical analyses that so many authors rely on them to 

corroborate other sources of evidence pertaining to past 

landscapes and events.

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Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 49

Oak Composition and Structure in the Eastern United States

W. Keith Moser1†, Mark Hansen2, Will McWilliams3, and Ray Sheffield4

Abstract.—Although oak species currently occupy a dominant position in most eastern deciduous forests, 

particularly on upland sites, many scientists and managers have expressed concern about the future of this 

genus in the absence of the disturbance patterns that facilitated its establishment up to now. Reductions 

in timber harvesting and fire in particular may give the advantage to competitors such as maples. Using 

data from the Forest Inventory and Analysis program of the U. S. Forest Service, we looked at current data 

and temporal trends to gauge the trajectory of oak forests in the Eastern United States. The area of the 

two upland oak groups—oak-hickory and oak-pine—covered 160.3 million acres or 43 percent of eastern 

timberland. The oak volume per acre of timberland has increased over the last four to five decades. Yet, we 

are seeing a decline in the proportion of total timberland with at least 20 ft2

ac-1 of select red or white oaks 

(the “select oak” stands). While the select oak basal-area component within these stands increased slightly, 

it represents a decreasing proportion of the total basal area in the stand, suggesting that associated species 

are increasing in their share of the overstory. While the total number of seedlings/saplings in the understory 

of stands with select red or white oak5 basal area greater than 20 ft2 ac-1 has been increasing, the proportion 

of all seedlings/saplings that are select white oak seedlings/saplings has been declining over the last 20 or 

so years. The declining proportion of regeneration represented by oak species suggests a future eastern U. 

S. forest with substantially reduced proportions of oaks in the overstory. Reintroducing disturbances such 

as fire is essential to maintain oaks’ overstory presence and associated biological and economic benefits.

that lead to eventual canopy occupancy (see Abrams, this 

volume); in this paper we examine trends, status, and 

implications of the structure and composition of oak 

forests in the Eastern United States.

We used data from the national forest inventory and 

analysis program (FIA) of the USDA Forest Service 

(Frayer and Furnival 1999). The FIA program conducts 

comprehensive forest inventories to estimate the area, 

volume, growth, and removal of forest resources in the 

United States, and measures the health and condition 

of these resources. The program’s sampling design has a 

base intensity of one plot per approximately 6,000 acres 

and is assumed to produce a random, equal probability 

sample. The national FIA program consists of five 

regional programs6 that provide estimates of forest 

area, volume, change, and forest health throughout the 

United States (McRoberts 1999). We used data from 

three of these regional FIA programs—North Central, 

Northeastern and Southern—to depict forest conditions 

for the Eastern United States. For historical data we used 

data generated from past Forest Inventory reports for 

Introduction and methodology

Oaks have been in eastern U. S. forests for at least 6,000 

years Lorimer (1993). While current oak forests evolved 

through a combination of ecological and humaninfluenced factors (McWilliams et al. 2002), changes 

in disturbance patterns are altering stand development 

trajectories to the detriment of oak (Larson and Johnson 

1998; Smith 2005). Other authors at this conference will 

present their interpretation of oak regeneration patterns 

5

The categories “select red oaks” and “select white oaks” have 

historically referred to those species preferred by mill owners 

for their uniform characteristics, quality and yield of higher 

grades of lumber. In examining trends over the last 25 or so 

years, we categorized stands as select red or white oak acreage 

as in those stands with at least 20 ft2ac-1 of the total basal 

area in select red white oak species and limited our analyses 

to those categories. These trends reflect management and 

utilization of those species that are most identified with the 

name “oak” and that comprise a significant proportion of the 

genus’ volume.

6

Soon to be four. The North Central and Northeastern FIA 

programs will merge into the Northern FIA program by 1

October, 2006.

1

USDA Forest Service, Northern Research Station, Forest 

Inventory and Analysis Program, 1992 Folwell Ave., St. 

Paul, MN 55108; 2USDA Forest Service, Northern Research 

Station, 1992 Folwell Ave., St. Paul, MN 55108; 3USDA 

Forest Service, Northern Research Station, Forest Inventory 

and Analysis Program, 11 Campus Blvd., Suite 200, Newtown 

Square, PA 19073; Southern Research Station, Asheville, NC; 

†

Corresponding author, 651-649-5155, email: wkmoser@

fs.fed.us.
50	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

states in the Eastern United States and data generated by 

the FIA Mapmaker program (Miles 2005). For current 

structure and regeneration, we used data generated by 

the FIA database.

Of the major deciduous forest-type groups in the 

Eastern United States, oaks are common associates 

of all but the northernmost groups of spruce-fir and 

aspen birch. Oak is most prevalent in the two upland 

oak groups: oak-hickory and oak-pine. Oaks also are 

members of other upland deciduous forest types. As 

defined by the FIA, upland oak forest type groups 

included eight detailed types within the oak-pine 

group and 11 oak types among the 17 types within the 

oak-hickory group. These groups are defined by the 

proportion of total stocking represented by oaks. Oakhickory group includes stands where half or more of the 

stocking is contributed by oak or oak-dominated stands. 

For the oak-pine group, stocking of oaks and other 

deciduous species is from 25 to 50 percent (McWilliams 

et al. 2002).

Results

Current Distribution of Mixed-oak 

Stands in Eastern United States

We used inventory data from the FIA database to 

identify the presence of oaks in the Eastern United 

States. Figure 1 is a map that displays the proportion 

of the total basal area occupied by all oak species. The 

occurrence of oak throughout forest stands in the east is 

readily apparent. This map shows that while eastern oaks 

are present from Maine to Louisiana and Minnesota to 

Florida, they comprise the most dominant portion of the 

canopy in the Ozarks and in portions of the Appalachian 

Mountains. Other areas with a high proportion of 

oaks include the Central Lowlands of Minnesota and 

Wisconsin, and the Tennessee River valley.

The area of the two upland oak groups covered 160.3 

million acres or 43 percent of eastern timberland in the 

most recent inventories (Table 1). Upland oak forests 

compose at least 48 percent of the timberland in the 

Figure 1.—Oak species’ percentage of total timberland basal area, based on the most recent 

FIA inventories from each state.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 51

Table 1.—Area of timberland in the Eastern United States, in millions of acres and from the most recent inventory, by region and broad forest type, with percent of region total11Source: Forest Inventory Database (Miles 2005). aIncludes oak-hickory and oak-pine forest-type groups.bIncludes swamp chestnut oak - cherrybark oak, sweetgum-Nuttall oak-willow oak, and overcup oak-water hickory forest types. cMichigan, Minnesota, and Wisconsin.dIllinois, Indiana, Iowa, Missouri.eConnecticut, Massachusetts, Maine, New Hampshire, Rhode Island, and Vermont.fDelaware, Maryland, New Jersey, New York, Ohio, Pennsylvania, and West Virginia. gFlorida, Georgia, Kentucky, North Carolina, South Carolina, and Virginia.hAlabama, Arkansas, Louisiana, Mississippi, Oklahoma, Tennessee, and Texas.iKansas, Nebraska, North Dakota, and South Dakota.

Total Upland oaka

Other upland 

deciduous Lowland oakb

Other lowland 

deciduous Conifer Other

Area Percent Area Percent Area Percent Area Percent Area Percent Area Percent

Lake Statesc 49.08 8.30 17 24.90 51 - 0 3.97 8% 11.39 23 0.52 1Central Statesd 25.21 19.06 76 2.02 8 0.09 0 2.96 12 0.85 3 0.23 1New England Statese 31.31 4.52 14 16.58 53 - 0 0.81 3 9.30 30 0.10 0Mid-Atlantic Statesf 56.03 26.93 48 22.50 40 0.11 0 2.21 4 3.82 7 0.46 1Atlantic statesg 96.01 47.84 50 1.04 1 3.91 4 9.60 10 32.99 34 0.62 1Gulf statesh 105.85 52.05 49 0.10 0 8.03 8 10.13 10 35.00 33 0.53 1Plains statesi 5.33 1.56 29 0.55 10 0.01 0 1.32 25 1.67 31 0.23 4Total 368.82 160.27 43 67.7 18 12.15 3 30.98 8 95.03 26 2.69 1
5	 Fire 2 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Atlantic, Central, Gulf and Mid-Atlantic states. Other 

upland deciduous groups cover 67.7 million acres or 

18 percent of eastern timberland. The Gulf states had 

the most oak, with 32 percent of the total eastern U.S. 

upland oak timberland area and 66 percent of the 

lowland oak timberland.

Oak volume has generally increased across the region 

(Fig. 2), in some cases at a declining rate. In upland 

oak stands with at least 20 ft2

 ac-1 of oak basal area, 

most acreage was in the sawtimber-size stands (Table 

2). There are several explanations for this distribution. 

First, oaks are long-lived species and thus spend a lower 

proportion of their total life in the seedling-sapling 

or poletimber size classes. Second, according to FIA 

protocol, mixed-age stands often are assigned to the size 

class of the largest component. For example, a stand that 

has one-third of the stocking in each size class is called 

sawtimber.

Some authors have suggested that the age-class 

distribution of oak stands is unbalanced and skewed 

toward older stands (Abrams and Nowacki 1992; 

Abrams 2005; Healy et al. 1997; Lorimer 1993). The 

FIA stand-size variable can provide some indication 

of the stages of stand development (Oliver and Larson 

1996), but the correlation with stand or tree age is less 

robust, because the classification is based solely on tree 

diameter (McWilliams et al. 2002). Each FIA plot has 

several “age” trees that are used to develop productivity 

equations. Because only the most dominant overstory 

trees are sampled, the ages may not represent all plot 

trees; those data are not considered here.

FIA has timberland area delineated by forest types 

which, in turn, are combined into forest type groups. We 

examined the components of upland oak type foresttype groups: oak-pine and oak hickory by forest type and 

stand size class (Table 2). The white oak/red oak/hickory 

Figure 2.—Average volume of oak species per acre of timberland (ft3 ac-1) by FIA program. North 

Central states: Illinois, Indiana, Iowa, Kansas, Michigan, Minnesota, Missouri, Nebraska, North 

Dakota, South Dakota, and Wisconsin; Northeastern states: Connecticut, Delaware, Maryland, 

Maine, Massachusetts, New Hampshire, New Jersey, New York, Ohio, Pennsylvania, Rhode Island, 

Vermont, and West Virginia; Southern states: Alabama, Arkansas, Florida, Georgia, Kentucky, 

Louisiana, Mississippi, North Carolina, Oklahoma, South Carolina, Tennessee, Texas, and Virginia.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 53

Table 2.—Timberland area of upland oak forest type groups by forest type and stand size class, for the states east of the Great Plains

Forest type Total Seedling-sapling Poletimber Sawtimber Nonstocked

Area

Percent of 

Total Area

Percent of 

Total Area

Percent of 

Total Area

Percent of 

Total

Oak/pine group 6.9 0.7 10 4.3

6

2 1.9

28 - - 

White pine/red oak/white ash 3,836.90

4

20.6 11 1,175.70 31 2,240.60 58 - - 

Eastern redcedar/hardwood

2,895.70 769.

2

2

7 1,481.50 51 644.9 22 - - 

Longleaf pine/oak 1,248.90 639.6 51 330.7

2

6

275.9 22

2.7

0

Shortleaf pine/oak 4,398.

2

0 579.9 13 1,438.80 33

2,379.50 54 - - 

Virginia pine/southern red oak

2,178.90 439.1 2

0 820.5 38 919.3

4

2 - - 

Loblolly pine/hardwood 15,79

2.10 6,104.70 39 3,722.50

2

4 5,959.80 38 5.1 0

Slash pine/hardwood 1,797.30 697 39 441.7

25 653.5 36 5 0

Other pine/hardwood

2,846.60 707.

2

25 978.7 34 1,157.80

41 2.9

0

Total oak/pine group 35,001.50 10,358.00 30 10,394.40 30 14,

233.

2

0 40 15.70

0

Oak/hickory group 1,444.10 922.5 64

243.7 17

277.9 19 - - 

Post oak/blackjack oak 5,680.30 808.5 14

2,460.90 43

2,410.90

4

2 - - 

Chestnut oak 5,7

28.70 165.4 3 1,750.10 31 3,813.

2

0 67 - - 

White oak/red oak/hickory 49,419.70 5,586.10 11 13,9

27.10

28 29,906.40

61 - - 

White oak 4,779.10

7

2.6

2 1,183.10

25 3,5

23.50 74 - - 

Northern red oak 3,78

2.2

0 110.5 3 805.5 21

2,866.

2

0 76 - - 

Yellow-poplar/white oak/red oak 7,450.90 739.7 10 1,674.50 22 5,036.60 68 - - Sassafras/persimmon 965

299.3 31 390 40

275.7

29 - - 

Sweetgum/yellow-poplar 6,649.

2

0

2,614.80 39 1,798.60

2

7

2,235.80 34 - - 

Bur oak 58

2.1 26.7 5 118.

2

2

0 437.

2 75 - - 

Scarlet oak 527.6 30.9

6

207.9 39

288.8 55 - - 

Yellow-poplar 1,250.

2

0 176.5 14 378 30 695.6 56 - - 

Black walnut 653.8 81.5 12 186.5

29 385.9 59 - - 

Black locust 474.9 168.

2 35

206.9 44 99.8 21 - - 

Southern scrub oak 1,281.10 904.8

71 326.6

25 39.5 3 10.

2 1

Chestnut oak/black oak/scarlet oak 3,051.70 177

6 944.4 31 1,930.40 63 - - 

Red maple/oak

2,381.60 368 15 965.1 41 1,048.50 44 - - 

Mixed upland hardwoods

29,166.60 7,640.70

2

6 8,851.60 30 12,65

2.10 43 22.2

0

Total oak / hickory group 125,

268.80

20,893.70 17 36,418.70

29 67,9

24.00 54 32.40

0

Total (Thousand acres) 160,

270.30 31,251.70 19 46,813.10

29 82,157.

2

0 51 48.1 0

Total(Thousand hectares) 64,861.40 12,647.60 18,945.30 33,

249.00 19.5
54	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

forest type was by far the largest with approximately 50 

million acres. The next largest forest type was in mixed 

upland hardwoods, with slightly less than 30 million 

acres. The loblolly pine/hardwood forest type was the 

third largest at around 15 million acres.

Using the most recent data, oak timberland area had a 

stand-size distribution of 19 percent seedling-sapling, 29 

percent poletimber and 51 percent sawtimber. The oak/

hickory forest type group summary was similar at 17, 

29, and 54 percent, respectively. The largest component 

of this group was the white oak/red oak/hickory, a forest 

type common in parts of the Southern Appalachians 

and the Ozark Plateau (McWilliams 2002). Sawtimber 

accounted for the majority of the acreage in this forest 

type, with 61 percent of the area classified as sawtimbersized stands and only 11 percent classified as seedlingsapling. Other individual oak-hickory forest types with 

percentages of the total area in seedling/saplings that 

were less than the oak/hickory group as a whole included 

chestnut oak (3 percent), post oak/blackjack oak (14 

percent), white oak (2 percent), northern red oak (3 

percent), bur oak (5 percent) and scarlet oak (6 percent).

The oak-pine forest type group had a more balanced 

stand structure, with 30 percent of the area in seedlingsaplings, 30 percent in poletimber, and 40 percent in 

sawtimber. This group’s largest component, loblolly 

pine/hardwood, distributed 39 percent seedling-sapling, 

24 percent poletimber, and 38 percent sawtimber. The 

forest types with percentages of the total area in seedling/

saplings that were less than the oak/pine group as a 

whole included white pine/red oak/white ash with 11

percent seedling-sapling, shortleaf pine/oak 1(3 percent) 

and Virginia pine/southern red oak (20 percent).

It is interesting to note both the higher average 

percentage of seedling-sapling timberland area in 

oak-pine forests vs. oak-hickory forests and the greater 

percentage of regeneration among those oak-pine types 

that are considered below the forest-type group average 

compared to oak-hickory below-average forest types. 

McWilliams et al. (2002) stated that these young oakpine stands commonly convert to pine stands as the pine 

species outgrow their competitors, so how much of these 

oak-pine seedling-sapling areas will result in mature 

forests with a significant component of oak remains to 

be seen.

Oak Overstory: Status And Trends

To examine overall trends in oak forest species in 

the overstory, we divided the estimated eastern U.S. 

timberland into three categories based on FIA plot-level 

data: stands with oak basal area greater than or equal to 

20 ft2

ac-1 (OAK 20 PLUS), stands with oak basal area 

less than that amount but greater than zero (OAK LT 

20), and stands with no oak basal area (OAK ZERO) 

(Fig. 3). The results vary across the region, but there is a 

slight decline in the percentage of plots in the “oak basal 

area greater than or equal to 20 ft2

ac-1” category and the 

proportion of plots with zero oak basal area is increasing. 

We can conclude that there is a downward trend in oak 

basal area but as yet no precipitous decline across the 

region.

We also examined upland oak overstory growing-stock 

volume according to individual species. Table 3 shows 

the volumes in descending order for species in the red 

and white oak groups. Northern red oak had the largest 

red oak volume followed by black oak. White oak had 

the largest white oak group volume followed by chestnut 

oak.

Mortality estimates are the principal indicator of health 

from FIA inventories and are calculated on an equivalent 

annual basis (McWilliams et al. 2002). Overall oak 

mortality was 0.79 percent (Table 3). The mortality of 

white oak species averaged 0.55 percent while red oak 

species’ mortality averaged 0.98 percent of growingstock volume. Those red oak species with the highest 

mortality included pin oak (1.78 percent), Nuttall oak 

(1.74 percent) and scarlet oak (1.34 percent). The white 

oak group species with the highest mortality rate was 

overcup oak at 0.81 percent.

McWilliams et al. (2002) noted that species with the 

highest mortality are subjected to the most stress agents 

and gave examples of American elm and Dutch elm 

disease and balsam fir and spruce budworm. While there 

could be a host of localized factors affecting mortality, 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 55

Figure 3.—Percentage of timberland area by oak category class, state and decade of inventory. Multiple 

inventories in one decade are averaged. “No oak” = 0 ft2 ac-1 oak species basal area; “Oak 0-20” = 0-19.9 ft2

ac-1 oak basal area; “Oak 20-plus” = 20+ ft2 ac-1 oak species basal area.

disease situations that are density- and age-mediated, 

such as oak decline in Missouri (Lawrence et al. 2002), 

could help explain the apparent high mortality of scarlet 

oak and pin oak.

Net Growth and Removals

The presence of a particular species is influenced not 

only by environmental considerations but also by how 

human activity impacts the species. A useful indicator 

of oak resource dynamics (McWilliams et al. 2002) is 

the ratio of growth to removals. Ratios less than 1.0 

indicate overcutting of the resource while values above 

1.0 indicate inventory expansion. We examined the 

latest estimates of net volume growth and removal 

volume by species (Table 4). The ratio for all oak species 

was positive; species that were most closely matched in 

terms of growth and removals included southern red 

oak (1.05), Nuttall oak (0.88), willow oak (1.17), and 

scarlet oak (1.18), suggesting that utilization of the oak 

resource is regionally sustainable. The high proportion 

of sawtimber stands implies that net growth will be 

primarily from increases in current tree diameter rather 

than ingrowth (McWilliams et al. 2002).

Status of Proportion of Oaks in 

Eastern U.S. Forest Overstory

In most of those states with select red oak (Table 4) 

stands that met the previously-mentioned criteria, mean 

overstory basal area of select red oaks increased from 

the 1980’s through 2003. Because oak regeneration 

potential is at least partially related to the proportion of 

oaks in the overstory (Johnson et al. 2002), it would be 

helpful to examine trends in the proportion of oaks in 

the total stand basal area. As the underlying theme of 

this paper is “where oak is going,” we examined trends 

in the percentage of total basal area that is in select red 

oaks (Fig. 4). We looked at changes between inventories 

in the 1980’s vs. 1990’s, 1980’s vs. 2000’s, and 1990’s 

vs. 2000’s. While some states showed an increase in the 

proportion of overstory basal area represented by SRO, 

most states showed an overall decline during this period.

The situation was similar for white oaks. Regionwide, 

there was an increase in white oak basal area in stands 

with at least 20 ft2

ac-1 of white oak, but this basal area 

represents a decreasing proportion of total stand basal 

area (Fig. 5).
56	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Table 3.—Growing-stock volume, mortality, growth and removals, in cubic feet, of oak species groups in 

the Eastern United States (excluding the Plains states); data based on the latest inventory for each state

Species

Growing-stock 

volume Mortality

% 

Mortality Net growth Removals

Growth 

removals ratio

Red Oak

 northern red oak 20,655,454,559 153,296,330 0.74 538,477,979 329,792,788 1.63

 black oak 12,630,532,249 153,531,016 1.22 332,118,409 231,452,931 1.43

 water oak 8,018,169,993 83,017,464 1.04 344,814,570 257,593,597 1.34

 southern red oak 6,535,917,882 56,645,858 0.87 235,162,961 224,973,434 1.05

 scarlet oak 6,460,923,509 86,851,528 1.34 157,857,759 134,157,620 1.18

 laurel oak 3,374,331,616 35,516,992 1.05 137,585,927 99,398,151 1.38

 willow oak 3,018,642,800 35,342,884 1.17 112,112,411 95,500,704 1.17

 cherrybark oak 2,695,838,154 18,276,233 0.68 114,284,906 91,056,971 1.26

 pin oak 804,771,728 14,337,991 1.78 21,554,964 15,717,424 1.37

 northern pin oak 709,967,448 4,309,812 0.61 21,454,369 6,079,116 3.53

 Nuttall oak 634,755,257 11,015,356 1.74 12,282,211 13,956,389 0.88

 Shumard oak 557,716,992 3,466,047 0.62 23,187,254 16,453,650 1.41

 live oak 513,642,097 1,082,584 0.21 11,480,662 3,167,426 3.62

 shingle oak 402,528,673 2,815,132 0.70 17,596,495 2,769,956 6.35

 blackjack oak 280,545,827 2,851,693 1.02 10,777,416 722,612 14.91

Total red oak 67,293,738,784 662,356,920 0.98 2,090,748,293 1,522,792,769 1.37

White Oak

 white oak 28,401,475,433 139,793,091 0.49 843,535,496 568,816,692 1.48

 chestnut oak 12,831,623,025 96,296,070 0.75 259,564,479 160,100,511 1.62

 post oak 6,273,795,886 33,244,411 0.53 212,414,329 136,672,484 1.55

 bur oak 1,936,207,204 4,064,910 0.21 93,947,682 15,584,760 6.03

 overcup oak 1,522,137,119 12,326,322 0.81 40,340,051 29,043,911 1.39

 chinkapin oak 904,100,004 2,287,263 0.25 28,954,892 6,237,172 4.64

 swamp chestnut oak 883,000,453 5,083,805 0.58 24,765,460 20,524,158 1.21

 swamp white oak 417,660,508 1,159,890 0.28 14,165,984 4,016,859 3.53

 Durand oak 24,008,130 0.00 470,805 --

 Delta post oak 20,239,202 0.00 833,243 514,152 1.62

 dwarf post oak 7,357,431 0.00 674,570 --

Total white oak 53,221,604,395 294,255,762 0.55 1,519,666,991 941,510,699 1.61

Total oak 120,515,343,177 956,612,682 0.79 3,610,415,284 2,464,303,469 1.47

and Johnson 1998; Johnson et al. 2002; Van Lear and 

Waldrop 1989; Van Lear 1991). However, human 

land-use changes altered disturbance type and intensity 

(Abrams 2005; Smith 2005), creating environmental 

conditions more favorable to other species (Parker and 

Merritt 1995; Larsen and Johnson 1998; Smith 2005.

Table 4.—Select red and white oak species

Select red oaks Select white oaks

Northern red oak White oak Chinkapin

Cherrybark oak Swamp white oak Durand oak

Shumard oak Swamp chestnut oak Bur oak

Oak Past

Historical disturbance patterns such as clearcuts, fire, or 

land clearing contributed to oak’s current dominance 

in upland forests (Liptzin and Ashton 1999; Rogers 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 57

Figure 4.—Percent change in proportion of total basal area of select red oak species 

in stands with at least 20 ft2 ac-1 of select red oaks, by state and inventory decade.

Figure 5.—Percent change in the proportion of total basal area of select white oak species in 

stands with at least 20 ft2 ac-1 of select white oaks, by state and inventory decade.
58	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

We examined temporal trends in the 

oak resource as a component of total 

timberland acreage. The dynamic 

nature of property ownership in this 

region affects the amount of land 

area in the timberland base and these 

changes impact all species. Timberland 

acreage was highest in the early decades 

followed by a gradual decline to a low 

in the 1970’s that was sparked by the 

intensification of land use, particularly 

the conversion to agriculture in the 

50’s and 60’s, the decline of open 

space, particularly forestland, to urban 

development (See Smith, this volume). 

From 1970 to present, timberland 

area increased due to the gradual 

abandonment and reforestation of 

farmland, particularly during periods of 

economic downturn in the farm sector.

Oak Future

A common technique for regenerating oaks is the 

two-cut shelterwood method (Johnson et al. 2002). 

To take advantage of oak’s life history strategy, the 

first cut removes enough overstory to create growing 

space (light on the forest floor) for regeneration. While 

seedlings/saplings may not grow rapidly in this partial 

tree shade in the understory (Johnson 1993; Larsen and 

Johnson 1998), the seedlings/saplings will maintain 

and expand a root system even while being topkilled 

repeatedly. With the established root systems, many 

of the seedlings/saplings are able to grow faster after 

removal of the residual overstory during the final cut of 

the shelterwood.

This silvicultural treatment mimics the perceived natural 

disturbance regime, exemplified by fire, that facilitated 

natural oak establishment in eastern hardwood stands 

(Johnson et al. 2002). Oak seedlings/saplings become 

established in the understory, sometimes in gaps 

(Ashton and Larson 1996; Rentch et al. 2003) and then 

undergo a series of topgrowth and dieback until some 

disturbance releases them to grow to the canopy.

Regeneration is heavily influenced by the size and 

composition of the forest overstory (Smith et al. 1996). 

We have seen a declining trend in the proportion of total 

timberland in oak overstory, with several exceptions, 

throughout the Eastern United States. Given the 

relatively long lives of certain oak species, the current 

oak presence likely reflects disturbance conditions far in 

the past. However, oak regeneration, should reflect more 

recent disturbances (or the lack of them). Accordingly, 

we looked at oak seedling/sapling data from the last one 

to three inventories in each state to gain insight into the 

future of oak forests.

The Eastern U.S. forests are not lacking for regeneration. 

In most states, we have observed increasing seedling/

sapling densities of all species between the 1980’s and 

2000’s (Fig. 6). Observed declines in seedling/sapling 

numbers occurred in certain states between inventories 

in the 1990s and in the 2000s, and may merely reflect 

the severe drought conditions in the Midwest (Lawrence 

et al. 2002) rather than a long-term trend.

Red Oak Regeneration

Figure 6.—Percentage change in trees per acre of all species of 

seedlings/saplings on all timberland, between inventory decades.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 59

We first looked at seedlings/saplings of 

select red oak species in what we defined 

as select red oak stands. Of these species, 

northern red oak is the most important 

component of this category and 

apparently benefited from the large-scale 

anthropogenic influences of the 19th and 

20th centuries (Abrams 2005) and other 

factors. Across the Eastern United States 

we observed increases or slight decreases 

in the number of select red oak seedlings/

saplings per acre over time (Fig. 7). While 

several states showed dramatic gains in 

the number of seedlings, most of the 

changes were more modest.

As with the analysis of red oak in the 

overstory, we were interested in temporal 

trends in the proportion of red oak 

seedlings/saplings to the total number 

of seedlings/saplings. Across most of 

the states, we have observed a decline in 

the percentage of all seedlings/saplings 

represented by select red oak species over 

time (Fig. 8).

White Oak Regeneration

As with the red oak seedlings/saplings, 

we have white oak regeneration 

information from the last 20 or so years, 

with some states better represented than 

others. White oaks were an extremely 

important component of the presettlement landscape (Abrams 2005) 

and still have the greatest growing-stock 

volume of any oak species (Table 4). 

White oaks are more shade tolerant than 

red oaks, so given the increasing density of 

deciduous forests in the eastern U.S., we thought there 

might be evidence of seedling/sapling accumulation in 

the understory.

While the total number of seedlings/saplings in the 

understory of stands with select white oak basal 

area greater than 20 ft2

 ac-1 has been increasing, the 

Figure 7.—Select red oak seedlings/saplings per acre, by state and 

inventory year.

Figure 8.—Percent change in the proportion of all seedlings/saplings that are 

select red oak species between decades, in stands with at least 20 ft2 ac-1 of 

select red oak overstory. For example, “1980 – 1990” compares an inventory 

conducted in the 1990’s with the same state’s inventory conducted in the 

1980’s.

proportion of all seedlings/saplings that are select white 

oak seedlings/saplings has been declining over the last 

several inventories (Fig. 9). The decline in white oak 

proportion, extending across the entire eastern United 

States and over several decades, suggests that this trend 

is neither temporary nor an anomaly of the data, but 
60	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

a representation of an established 

trajectory in oak regeneration.

Summary

After reaching its low point in the 1960’s 

and 1970’s, timberland in the eastern 

U.S. has recently started to increase. 

The oak volume per acre of timberland 

has increased over the last four to five 

decades as overstory trees increased in 

size. While total oak volume has been 

increasing, there is a decline in the 

proportion of total timberland with at 

least 20 ft2

ac-1 of select red or white oaks 

(the “select oak” stands). Within these 

stands, the select red or white oak basalarea component increased slightly. While 

the oak basal area has been increasing, it 

represents a decreasing proportion of the 

total basal area in the stand.

Johnson et al. (2002) detailed the importance of 

accumulating oak regeneration in the understory and 

outlines the disturbances, anthropogenic and natural, 

that encourage this accumulation. Such disturbances 

promote two processes: the concentration of early 

growth on the oak seedling/sapling root system resulting 

from repeated dieback of the above-ground component, 

and the elimination of less fire-resistant species that 

otherwise would compete successfully for resources. Two 

of the most prominent disturbances are harvesting and 

fire. Both disturbances have been declining in eastern 

oak forests in recent times. Contributing to decreasing 

proportions of oak seedlings/saplings and seedlings/

saplings as reflected in our data across the eastern United 

States.

The declining proportion of regeneration represented 

by oak species is particularly disquieting as it provides a 

foretelling of future forest overstory composition. Given 

these regeneration trends, it is hard to imagine a future 

eastern U.S. forest landscape with the current proportion 

of oaks in the overstory. While reinstituting disturbances 

such as fire will be problematic in the highly populated 

landscape of this region, it should be considered part of 

the toolbox that resource managers use as they seek to 

maintain the many benefits that oak forest provides.

Literature cited

Abrams, M.D. 2005. Ecological and ecophysiological 

adaptations and responses to fire in eastern oak 

forests. In: Fire in eastern oak forests: delivering 

science to land managers, conference proceedings; 

2005 November 14-16.

Abrams, M.D.; Nowacki, G.J. 1992. Historical 

variation in fire, oak recruitment, and postlogging accelerated succession in central 

Pennsylvania. Bulletin of the Torrey Botanical Club. 

119: 19-28.

Ashton, P.M.S.; Larson, B. C. 1996. Germination 

and seedling growth of Quercus (section 

Erythrobalanus) across openings in a mixeddeciduous forest of southern New England, USA.

Forest Ecology and Management. 80: 81-94.

Frayer, W.E., Furnival, G.M. 1999. Forest survey 

sampling designs: A history. Journal of Forestry. 

4-10.

Figure 9.—Percent change in the proportion of all seedlings/saplings that are 

select white oak species between decades, in stands with at least 20 ft2 ac-1 of 

select white oak overstory.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 61

Healy, W.M.; Gottschalk, K.W.; Long, R.P.; Wargo, 

P.M. 1997. Changes in eastern forests: chestnut is 

gone. Are oaks far behind? Transactions of North 

American Wildlife and Natural Resources Conference. 

62: 249-263.

Johnson, P.S.; Shifley, S.R.; Rogers, R. 2002. The 

ecology and silviculture of oaks. New York: CABI 

Publishing. 501 p.

Johnson, P.S. 1993. Sources of oak reproduction.

In: Oak regeneration: serious problems, practical 

recommendations. Loftis, D.L.; McGee, C.E., 

eds. Gen. Tech. Rep. SE-84. Asheville, NE: 

U.S. Department of Agriculture, Forest Service, 

Southeastern Forest Experiment Station: 112-131.

Larsen, D.R.; Johnson, P.S. 1998. Linking the ecology 

of natural oak regeneration to silviculture. Forest 

Ecology and Management. 106(1): 1-7.

Lawrence, R.; Moltzan, B.; Moser, W.K. 2002. Oak 

decline and the future of Missouri’s Forests.

Missouri Conservationist. 63(7): 11-8.

Liptzin, D.; Ashton, P.M.S. 1999. Early-successional 

dynamics of single-aged mixed hardwood stands 

in a southern New England forest, USA. Forest 

Ecology Management 116: 141-150.

Lorimer, C.G. 1993. Causes of the oak regeneration 

problem. In: Oak regeneration: serious problems, 

practical recommendations. Loftis, D.L.; McGee, 

C.E., eds. Asheville, NC: Gen. Tech. Rep. SE-84. 

U.S. Department of Agriculture, Forest Service, 

Southeastern Forest Experiment Station: 14-39.

McRoberts, R.E. 1999. Joint annual forest inventory 

and monitoring symposium: The North Central 

perspective. Journal of Forestry. 27-31.

McWilliams, W.H.; O’Brien, R.A; Reese, G.C.; Waddell, 

K. L. 2002. Distribution and abundance of oaks in 

North America. In: McShea, W.J.;Healy, W.H., eds. 

Oak forest ecosystems: ecology and management for 

wildlife. Baltimore, MD: Johns Hopkins Press: 13-32.

Miles, Patrick D. 2005. Forest inventory mapmaker 

web-application version 2.1. St. Paul, MN: U.S. 

Department of Agriculture, Forest Service, North 

Central Research Station. www.ncrs2.fs.fed.us/4801/

fiadb/index.htm]

Oliver, C.D.; Larson, B.C. 1996. Forest stand dynamics 

2nd ed. New York: J Wiley.

Parker, G.R.; Merritt, C. 1995. The Central Region. In: 

Barrett, J.W., ed. Regional silviculture of the United 

States. New York: J. Wiley: 129-172.

Rentch, J.S.; Fajvan, M.A.; Hicks, Jr., R.R. 2003. Oak 

establishment and canopy accession strategies in 

five old-growth stands in the central hardwood 

forest region. Forest Ecology and Management. 184: 

285-297.

Rogers, R.; Johnson, P.S. 1998. Approaches to 

modeling natural regeneration in oak-dominated 

forests. Forest Ecology and Management. 106: 45-54.

Smith, D.M.; Larson, B.C.; M.J. Kelty, M.J.; Ashton, 

P.M.S. 1996. The Practice of silviculture, applied 

forest ecology, 9th ed. New York: J Wiley.

Smith, D.W. 2005. Why sustain oak forests? In: Fire 

in eastern oak forests: delivering science to land 

managers, conference proceedings; 2005 November 

14-16.

Van Lear, D.H. 1991. Fire and oak regeneration in the 

southern Appalachians. In: Nodvin, S.C.; Waldrop, 

T.A., eds. Fire and the environment: ecological and 

cultural perspectives. Gen. Tech. Rep.SE-69. Asheville, 

NC: U.S. Department of Agriculture, Forest Service, 

Southeastern Forest Experiment Station: 15-21.

Van Lear, D.H., Waldrop, T.A. 1989. History, use 

and effects of fire in the Appalachians. Gen. 

Tech. Rep. SE-54. Asheville, NC: U.S. Department 
6	 Fire 2 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

WHY SUSTAIN OAK FORESTS?

David Wm. Smith1

Abstract.—A brief overview and some personal thoughts are offered that deal with the implications of our 

social and political systems on the long-term sustainability of our forest resources. The connection of the 

most recent climatic events, in a geologic-time context, to the development of present day oak dominated 

forests of the Eastern United States is discussed. The impacts of human activity and human infrastructure 

during the recession of the Wisconsin Glacier that began about 15,000 years ago to the present are reviewed. 

Changes in eastern oak forests since European settlement in the early 1600s, and more specifically in the 

last half century are presented in greater detail. Also discussed are important characteristics of the more that 

30 oak species native to the Eastern United States, the complexity of the oak dominated eastern forests, the 

uniqueness of oak species for a variety of forest products and uses to satisfy human needs, and the critical 

importance of the oaks for wildlife food and cover. Finally, seven reasons for sustaining oak forests are presented.

scientific knowledge and professional experience, or 

when passage is influenced by issues and circumstances 

that are not related to the legislation, is partisan, or 

caters to special interests. Another concern is that 

there is an ever growing disconnect between what we 

use and where it comes from. This disconnect has 

worsened as we moved from a predominately rural 

to a predominately urban population over the past 

100 years. The concept that toilet paper comes in 

a roll and not from a tree and that milk comes in a 

container and not from a cow pasturing in a field has 

far reaching implications when it comes to the need 

to practice forestry that will provide people across the 

country and world with the products, and values and 

uses they depend on and demand. We also must deal 

with a general public that has little understanding 

of the dynamics of forest establishment, growth and 

development. For the majority of our society forests are 

viewed like a picture or a “snapshot in time”—a view of 

a forest as an entity that is static and does not change. 

The fact is that the only thing constant in a forest is 

change.

OAK FORESTS IN THE RECENT 

GEOLOGIC PAST

To gain ideas and a perspective on how to develop 

and organize this presentation, I talked to several 

people who were knowledgeable about and interested 

in eastern hardwoods and the implications of fire on 

forest stand development in oak and mixed-oak stands 

in particular. One person suggested that I explore what 

may be known about oaks and oak species evolution 

INITIAL THOUGHTS

How we will achieve sustainable management of our 

oak forests is complex and encompasses much more that 

just understanding the biological aspects of trees and 

forests. It has become evident how differently various 

groups look at or perceive forests. For most foresters 

and land owners who grew up on farms or have been 

intimately associated with their forested land, we think 

about the evolution and changes that occur over periods 

of tens and hundreds of years. We plan and carry out 

management activities, the full results of which we may 

not live to see. There is a significant body of scientific 

knowledge and professional experience that we use 

to make these long-term decisions. Is this knowledge 

complete? Certainly not, but it is sufficient for us to 

move forward and be relatively sure that we are moving 

toward our goal of sustaining our forested ecosystems. 

There are several obstacles that make the road to success 

a bit difficult. We live in a democratic country and 

our government functions through a well-established 

political system. It is through this system that all 

legislation (including that related to forests, forestry and 

the environment) is formulated, discussed, and enacted 

or rejected. With this political system there are some 

drawbacks. We often are faced with new legislation 

that results in what I term “political silviculture.” This 

is forest resource management policy legislation that 

often includes silvicultural practices or constraints on 

silvicultural practices that denies or disregards established 

1

Emeritus Professor of Forestry, Department of Forestry, 

Virginia Tech, Blacksburg, VA 24061, 540-552-6705, email: 

smithdwm@vt.edu
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 63

in terms of geologic time. I thought for a moment and 

the idea of tens of millions of years did not appear to 

be relevant to this particular conference. In fact, just 

thinking about what might have happened during the 

Pleistocene geologic epoch (1.8 million to 8000 years 

ago) seemed to be more than we really needed to be 

concerned about. When you consider that glaciers came 

southward an estimated 17 times during the Pleistocene 

and our eastern forests migrated north and south the 

same number of times (Goudie 1992), I do not think 

it is necessary to concern ourselves with what happened 

prior to that geologic period. What seems to be most 

important to the question of whether we should attempt 

to sustain the oak dominated forests of the Eastern 

United States is what has happened since the last ice 

age and what has been the influence of humans on the 

development of today’s forests.

The Wisconsin Glacier, the most recent glacial period 

of the Pleistocene, began about 100,000 years ago and 

reached its southern most extent in the Eastern United 

States about 15,000 years ago. (Nilsson 1983; Porter 

1988). If we were to look at the Eastern United States 

15,000 to 18,000 years ago, the sequence from north to 

south would have been ice sheets/glaciers, tundra and 

cold steppes, then white spruce forests, followed by jack 

pine forests, southern pine-oak forests, and finally sand 

dune-scrub communities in the lower half of Florida 

(Bonnicksen 2000). The spruce forests extended into 

North Carolina (Whitehead 1967; Wright 1981) and as 

far south as central Missouri, Arkansas, and southeastern 

Oklahoma. Some hardwoods including hornbeam 

and northern red oak may have been interspersed with 

the spruce in some places (Davis 1981; Delcourt and 

Delcourt 1991, 1979). A band of mixed hardwoods, 

including oak species, bordered the lower Mississippi 

River and major tributaries in protected areas adjacent to 

the spruce forests (Delcourt and Delcourt 1979, Wells 

1970). The jack pine forests were just to the south of 

the spruce forests and consisted of jack pine and red 

pine with a few temperate hardwood species scattered 

throughout the pine on favorable sites. The jack pine 

forests extended southward to central Georgia, Alabama, 

and Mississippi and westward to Memphis and the 

lower Mississippi River Valley (Delcourt and Delcourt 

1991; Whitehead 1973,;Wright 1981; Davis 1981). 

The mixed-hardwood, and southern pine-oak forests, 

lying to the south of the jack pine forests and containing 

most of the present day oak species, were relegated to the 

central and northern parts of Florida, and the southern 

parts of Georgia, Alabama, Mississippi, Louisiana, and 

Texas (Delcourt and Delcourt 1981; Porter 1988; Webb 

1987). The southern half of Florida was too dry and hot 

to support trees (Davis 1981; Wright 1981).

When considering this historical distribution of forests 

15,000 to 18,000 years ago, it is evident that those 

mixed hardwood, oak-pine, and pine forests occurred 

only in the southern portion of the country and 

occupied only about 20 percent of the land area they 

occupy today. In addition, the species composition, 

stand structure and distribution patterns probably were 

different from those of the present. Since the last glaciers 

started to recede, there has been a dramatic migration 

northward coupled with a rapid evolution of species 

composition and community structure, and adaptation 

to complex climatic and geographic changes. Since the 

end of the last Ice Age, the only thing constant in our 

Nation’s forests has been change. I think the following 

quote puts the evolution of our forests in perspective. 

“Forests only exist in human minds. Groups of animals 

and plants that we call forests come together for a short 

time; then each species goes its separate way when 

conditions change. Constant warming and cooling of 

the climate, and the ebb and flow of glaciers, caused 

the disassembly of old forests and the reassembly of 

new forests. Some species thrive in glacial cold while 

others do best in the warm periods between glaciations. 

So different forests dominated the land under different 

climates” (Bonnicksen 2000).

The Holocene geologic epoch that we are presently in, 

began 8,000 to 10,000 years ago (USGS 2005). It is 

during this warming period that the glaciers retreated, 

our eastern forests began their migration north, and 

the evolution of today’s forests began. Oaks have been a 

part of the forests throughout this evolutionary process. 

As an example, oak pollen from sediment core samples 

of Holocene deposits in Cliff Palace Pond, a small 

woodland pond, located below a sandstone ridge in the 

northeastern part of Jackson County in southeastern 

Kentucky, revealed the presence of oak pollen 9,500 
64	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

years ago (Ison 2000; Delcourt et. al. 1998). Oak pollen 

showed a general increase to about 4,800 years ago, then 

declined dramatically for about a thousand years and 

then began to increase about 3,000 years ago. The oak 

pollen has remained at its present high for most of the 

past 3,000 years, with pollen from American chestnut 

and pine also being significant (Delcourt et. al. 1998). 

Delcourt and others measured charcoal particles and the 

relative abundance of fire-tolerant and fire-intolerant 

trees and shrubs. This study and similar studies have 

proved to be extremely helpful in investigating the role of 

fire, both natural and anthropogenic, in the regeneration 

of oak species in the upland hardwood and mixed oakpine forests of the Eastern United States.

HUMAN MEDIATED CHANGE IN OAK

FORESTS

In a Geologic Time Context

The presence of humans and the impacts of man’s 

activities have resulted in forests that are different than 

those of the previous glacial cycles when forests migrated 

north and south as the climate changed. Humans, 

migrating eastward from Siberia, across Beringia (now 

lying below the Bering Strait) and into Alaska, have 

been present in North America for at least 15,000 years 

(Bonnicksen 2000; Fiedel 1987; Fagan 1991, 1987). It 

may have taken another 2,000 to 3,000 years for these 

Paleoindians to work their way south and east into 

what is now the Eastern United States. Paleoindians 

probably reached southeastern Wisconsin between 

13,400 and 12,300 years ago (Bonnicksen 2000; Hall 

1998). During the last 8,000 to 10,000 years, as the 

Wisconsin ice sheet retreated, human activity has had 

an ever increasing influence on how hardwood forests 

developed as they migrated north. The First Americans 

migrated south as the climate warmed. By 12,000 years 

ago, Paleoindians had settled in Florida and had moved 

as far west as St. Louis, Missouri (Canby 1979; Graham 

et. al. 1981). People were here and they were here to stay. 

For the next 11,600 years, until the arrival of Europeans 

in the early 1600s, fire was the primary anthropogenic 

tool for mediating change in eastern forests. We do 

not know how much change was directly or indirectly 

attributed to human caused fire, but we are confident 

that fire was used and for several thousand years. With 

the initial European settlements along the East Coast and 

the rapid migration west came land clearing for farming, 

the construction of towns and cities, and a rapid increase 

in the human population of a growing nation. The 

building of America had begun and the forests of the 

East were in rapid transition—not from the gradual 

climate changes measured in geologic times of tens of 

thousands of years but from human-induced changes in 

a matter of decades.

Since European Settlement

Think for a moment about some of the human-caused 

or mediated events of the past 300 years that have 

modified or permanently changed the oak dominated 

forest landscapes of the Eastern United States. Probably 

the first and most widespread was the conversion of 

millions of acres of forest land to pasture for domestic 

animals and the production of agricultural crops. 

The chestnut blight, a disease caused by the fungus 

Cryphonectria parasitica, virtually eliminated the 

American chestnut from oak-chestnut forests of the 

east—to the point where we changed the forest cover 

type name to oak-hickory. The exclusion of periodic 

understory fire in the past 100 years has had a significant 

negative influence on the recruitment of oak species 

and a positive influence on the recruitment of species 

e.g., maple, beech, and blackgum, in many central and 

eastern hardwood forests (Abrams 2000, 2005). One 

of the most interesting and perhaps a most significant 

event was related to the annual migration of the now 

extinct passenger pigeon. Little is known about the 

ecological impacts of the passenger pigeon and it is rarely 

mentioned in the context of oak forests and changes in 

these forests that must have occurred when one considers 

the almost unbelievable numbers that were present. 

Scott Weidensaul (1994) wrote: “In 1808, in Kentucky, 

ornithologist Alexander Wilson watched the sky blacken 

with birds—a flock he estimated at a mile wide that 

passed him for four hours. They were passenger pigeons, 

easily the most abundant species of bird in North 

America and likely the most numerous land bird in the 

world. Based on his observations, Wilson calculated 

the flock’s number at 2.25 billion birds. They, and 

their kin, were the single greatest living expression of 

the bounty of the wooded sea.” Weidensaul noted that 

the pigeons existed in a relatively few enormous flocks, 

with each flock numbering in the hundreds of millions 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 65

of birds. “Several billion birds feeding through a forest 

would reduce the mast available to other nut-eaters and 

would seriously reduce tree regeneration; the fractured, 

mangled trees, collapsed under the sheer weight of the 

roosting or nesting birds (to say nothing of the vast 

quantities of guano), would have provided a flush of 

sunlight and fertilizer for new plant growth, benefiting 

grazers and browsers like elk and deer.” Passenger 

pigeons were still nesting by the millions in the 1870s, 

by the turn of the century they were almost gone from 

the wild, and the last known passenger pigeon died in 

the Cincinnati Zoo on September 1, 1914 (Weidensaul 

1994). Think about this for a moment: a bird species 

composed of billions becoming extinct in less than 50 

years. It was during this same period that major forest 

harvests of mature and old-growth oak forest types were 

taking place within the natural range of the passenger 

pigeon in the Eastern United States. Here were two 

major landscape altering events occurring at the same 

time—and both closely linked to oak forests. What 

were the individual and the combined effects of these 

two agents of change on the oak forests of today? It is 

certainly worthy of some thoughtful conversation and 

contemplation. 

Eastern Oak Forests and Recent 

Trends

The total land area of the Eastern United States, which 

includes Minnesota, Iowa, Missouri, Oklahoma, Texas, 

and all states to the East, is 948 million acres, or 42

percent of the U.S. land area. There are 384 million 

acres of forest land in the East, or 52 percent of the total 

U.S. forest land area. Today, the eastern U.S. is 40.5 

percent forested. How has this forest land area changed 

over time? In 1630 it is estimated that the eastern 

U.S. was nearly 70 percent forested (Fig. 1). By 1907, 

primarily as a result of land clearing for agricultural 

crops and pasture, the forest-land base had declined 

to 39.5 percent (652 to 374 million acres) of the total 

land area. It is believed that this was the low point 

in forest coverage in the eastern U.S. since sometime 

during the early Holocene epoch when the ice sheets 

were receding—some 8,000 to 10,000 years ago. Since 

the first part of the 20th century, the eastern U.S. forestland base has remained remarkably constant at about 40 

percent (Fig. 1).

The forest-land base has remained essentially constant 

but what about forest types and species? When looking 

at land area trends of the oak type groups, both upland 

and bottomland, over the past 50 years, there appears 

to have been an increase in the acreage of upland types 

and a decrease in the bottomland types (USDA For. 

Serv. 2003). The area occupied by the oak-pine type 

group increased by 30 percent and the oak-hickory type 

increased by 23 percent, while the bottomland oakgum-cypress type declined by 17 percent (Table 1). The 

conversion of productive bottomland sites from forest 

to agriculture uses and the oak-gum-cypress types to 

pine plantations are significant factors in the decline in 

bottomland hardwood acreage. The increase in upland 

oak type groups likely is the result of the reversion of 

agricultural land back to forest cover, especially land that 

was cleared but proved to be marginal for agricultural 

Figure 1.—Trends in U.S. forest-land 

area 1630-2002.
66	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

uses. Overall, there was a 15-percent 

increase in the total acreage of the oak type 

groups in the past 50 years (Table 1). It is 

important to note that these forest-type 

groups are broad general classifications and 

species composition within a type group 

could vary by 15 to 20 percent or more. 

For example, the oak-hickory type group 

is defined as: “forests in which upland 

oak or hickory, singly or in combination, 

comprise a plurality of the stocking except 

where pines comprise 25 to 50 percent, in 

which case the stand is classified as oakpine. Common associates include yellowpoplar, elm, maple, and black walnut” (USDA For. Serv. 

2003). Therefore, these data tell us little about possible 

shifts in the oak component within these type groups.

An analysis of the hardwood resource data from the 

Forest Service’s Forest Inventory and Analysis unit 

over the past 40 years revealed some interesting and 

compelling trends that may be helpful in obtaining a 

better understanding of what has been happening in 

eastern U.S. oak forests. The net volume of all hardwood 

growing stock on timberland has shown a remarkable 

increase of nearly 70 percent between 1963 and 2002

(Table 2). A significant part of this impressive net 

volume increase probably is related to the skewed ageclass distribution in eastern hardwood forests. Major 

harvests of original forests occurred in the Northeast, 

Lake States, Appalachian Mountains, and in the South 

between 1850 and 1920 (Sedjo 1991) The result is a 

significant bulge in the 80- to 150-year age classes where 

forest stands are vigorous and mortality is relatively low. 

The rapid increases in net volume growth during the 

past 40 years probably will level off in the next 10 to 20 

years. Since 1963, the proportion of select white and red 

oak net growing stock volume has remained about the 

same at 16 percent. However, the proportion of growingstock volume of all other oaks has shown a decline of 16 

percent. By contrast, the proportion of soft maple net 

volume has increased by 60 percent and that of yellowpoplar has increased by 35 percent. The proportion of 

net growing-stock volume of “all other hardwoods” has 

shown a slight decline of less than 5 percent (Table 2). 

The proportion of hard maple and ash (both species were 

included in the “all other hardwood” category of Table 1) 

net growing-stock volume has also increased in the past 

40 years (USDA For. Serv. 2003). If we consider the oaks 

to be an important component of eastern U.S. hardwood 

forests, then the general proportional decline of oak net 

growing-stock volume and the proportional increase in 

volume of associated species in the relatively short period 

of 40 years should be of major concern.

THE OAKS

Species and Distribution

Quercus is the classical Latin name of the oaks and is 

thought to be of Celtic derivation meaning fine and 

tree (Little 1979). The oaks are the largest tree genus 

in the United States and arguably the most important. 

Their worth is enormous when you consider the large 

number of species, the vast potential for forest products, 

the importance for wildlife food, cover and habitat, 

their importance for aesthetics and scenic beauty, their 

functional use in urban forests, and probably most 

important, their ecological value as significant functional 

components of most forested ecosystems, especially in 

the Eastern United States. Little (1979) recognized 58 

native oak species in the United States and Canada, and 

about 10 native oak shrubs. Over half of the 58 native 

species are found in the Eastern United States. Of the 

24 oak species described in significant detail in Silvics 

of North America—Hardwood, Volume 2 (Burns and 

Honkala 1990), 20 are considered eastern species and 

4 are only found in the western United States. Smith 

(1993) reviewed some of the regeneration-related silvical 

characteristics of 31 oak species, 25 of which were 

Table 1.—Timberland area in the Eastern United Statesa of oak type 

groups, 2002, 1997, 1987, 1977, and 1953

OakpineOakhickoryOak-gumCypressOak types 

Year total

-------------------------million acres-------------------------

2002 32.6 124.3 29.8 186.7

1997 33.4 124.0 29.3 186.7

1987 31.1 117.0 28.0 176.1

1977 34.6 108.6 26.6 169.8

1953 25.0 101.3 35.7 162.0

a Includes Minnesota, Iowa, Missouri, Oklahoma, Texas, and all states to the 

East.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 67

native to the Eastern United States and 6 to the western 

states. It is interesting that most eastern oak species 

occur in association with a wide variety of overstory and 

understory species. The combination of individual oak 

species and associated overstory and understory species 

are dependent on specific site conditions, and one or 

more species of oak is adapted to virtually every forest 

site condition that is found in the Eastern United States. 

In fact, if you were to overlay the ranges of just five 

species, northern red oak, white oak, bur oak, overcup 

oak and live oak, you would cover all sites in the Eastern 

United States except for several counties in northern 

New York. The overstory and understory species greatly 

influence oak regeneration distribution and success, and 

oak growth and development throughout the rotation. 

By contrast, the oaks of the Western United States 

generally are smaller in stature and in the total area that 

they occupy, Western oak species generally are found 

on more arid sites and in clumps or small pure stands. 

Competition from grasses and shrubs and drought often 

are major factors in regeneration success and subsequent 

seedling development on many western sites.

In general, oak species native to the Eastern United 

States are strong, durable, and relatively long-lived 

compared to associated species. Ages of 150 to 300 

years for northern red oak, white oak, and chestnut 

oak are not uncommon, and ages up to 700 years have 

been recorded (Hora 1981). Very large oaks have been 

documented in the east. Probably the best example is a 

white oak that was cut in 1913 near Lead Mine, West 

Virginia. This giant was 13 feet in diameter 16 feet from 

the base and 10 feet in diameter 31 feet from the base 

(Clarkson 1964). That amounts to 19,200 board feet 

(International 1⁄4 inch log rule) or 23,100 board feet 

(Doyle log rule) of lumber from the first log.

Uses and Values

The oaks serve as a raw material for a host of forest 

products and as such have a significant impact on local 

and regional economies throughout the eastern United 

States. The wood from oak is noted for its beauty, 

durability, strength, and decay resistance. Just think for a 

moment about the many products from oak that we take 

for granted, but use virtually every day: fine furniture, 

flooring, paneling, structural timbers, pallets, charcoal, 

veneer, molding, railroad ties, boxes and crates, cabinets, 

cooperage, boat building materials, handles, musical 

instruments, architectural woodwork, fuelwood, wood 

composites, and mulch.

The oaks also are important in the urban forest 

where they provide shade, food, and cover for urban 

wildlife, protection from wind, visual buffer zones, and 

aesthetically pleasing landscape components in parks, 

along streets, in yards, and in public areas.

A shift in forest tree species composition from the 

oaks to species including the maples, yellow-poplar, 

blackgum or beech has serious implications for many 

wildlife species in eastern hardwood forests. Acorns (hard 

mast) from oaks represent a significant food source for 

numerous mammals and birds, especially for fall and 

winter diets when sufficient food is critical for survival 

Table 2.—Net volume of hardwood growing stock on timberland in the Eastern United Statesa of 

selected species, 2002, 1997, 1987, 1977, and 1963

Total, all Select white

hardwood oak and Other Soft Yellow- All other

Year species red oak oaksmaple poplar hardwoods

100 million ft3 -------------------------Percent of all hardwood species-------------------------

2002 327.7 15.9 18.4 10.2 6.9 48.6

1997 316.4 16.2 18.2 10.2 6.7 48.7

1987 281.4 16.0 19.9 9.2 5.8 49.1

1977 241.3 16.4 21.0 7.9 5.7 49.0

1963 193.6 15.8 22.0 6.5 5.1 50.6

a Includes Minnesota, Iowa, Missouri, Oklahoma, Texas, and all states to the East.
68	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

(Martin et al. 1961). Wildlife, including deer, bear, wild 

turkey, quail, grouse, squirrel, mice and many songbirds 

depend on acorns for part of their diet. Some examples 

of these songbirds include the red-bellied woodpecker, 

tufted titmouse, blue jay and white-breasted nuthatch 

(Martin et al. 1961; Smith and Scarlett 1987; Grubb and 

Pravosudov 1994). Martin and others (1961) noted that 

acorns rate at or near the top of the wildlife food list, and 

listed more that 90 species of wildlife that use acorns. 

Acorns are a highly digestible, high-energy, low-protein 

food for many wildlife species (Kirkpatrick and Pekins 

2002). Compared to many other seasonal food sources, 

acorns are relatively slow to decompose and therefore 

available for an extended period when other food sources 

are limited or absent. Acorns ripen and are dispersed in 

the fall and are available throughout the winter while 

the seeds of red maple and silver maple ripen and are 

dispersed in late spring and early summer. The seeds of 

sugar maple and yellow-poplar ripen and are dispersed in 

the fall.

The annual production of acorn crops is highly variable 

and unpredictable and varies by species, region, and 

environmental conditions (Koenig and Knops 2002). 

Since hard mast often is a critical source of wildlife food, 

the variability in annual acorn production is a factor 

in limiting wildlife populations and greatly influences 

periodic population fluctuations. The consequences 

of declining oak species composition has serious 

ecological implications. The results of a recent study in 

Pennsylvania (Rodewald and Abrams 2002) is perhaps 

the first study to suggest that shifts in forest composition 

from oak toward maples may reduce bird species richness 

and abundance within forest bird communities and 

have a negative influence on certain species. The most 

likely species to be impacted are long-distance migrants, 

residents and bark-gleaning species. Few birds consume 

maple seeds (Martin et al. 1961). When the food value, 

quantity (weight), season of dispersal (soft maples), and 

decomposition rates of maple seed are compared with 

those of acorns, the latter clearly are superior.

Other physical factors of oaks versus maples and other 

species, such as bark thickness, maximum tree size, and 

longevity, also are important when evaluating shifts 

in species composition. The rough and deep furrowed 

nature of most oaks compared to the relatively smooth 

bark of red maple is an important and distinct advantage 

for insect-foraging bird species (Jackson 1970). The 

generally greater tree size, longer life, and the decay 

resistance of the oaks are important in their enhanced 

value as den and cavity trees. The larger size makes them 

more adaptable for bear and larger animals, and the fact 

that they live longer and are more resistance to wind fall 

and other climatic events adds to the stability of wildlife 

species that are dependent solely or in part on den cavity 

trees.

For a more comprehensive discussion of the importance 

of oak ecosystems to wildlife and the interactions of a 

host of wildlife species with these oak dominated forests 

see Oak Forest Ecosystems: Ecology and Management for 

Wildlife (McShea and Healy 2002).

WHY SHOULD WE SUSTAIN OAK

FORESTS?

There is little question that change is the only thing that 

is constant in forests. I believe that one of the overriding 

issues is not the fact that changes in oak composition and 

distribution are probably occurring all across the eastern 

forests but that the rapid rate of change is unprecedented 

in the context of biologic and ecologic time frames of 

the past. We are seeing changes in less than a single 

generation of oaks that we would have expected in 

10 to 20 or more generations. Ecological balances 

are not being reached as a result of human induced 

changes. It is imperative that we strive to enhance our 

understanding of the short and long-term, direct and 

indirect implications of human activity on our forested 

ecosystems. We must identify the information needed to 

understand the biological processes and functions that 

will result in forest resource management decisions that 

will ensure the future biological quality and ecological 

integrity of the forested-land base. We must diligently 

identify, prioritize, plan, conduct, and analyze, research 

programs that will fill the gaps in our present knowledge. 

Above all, we must report research results and translate 

and communicate them to people on the ground who 

are working to solve real-time problems. Finally, we must 

ensure that we have a built-in evaluation and follow-up 

system so that future information needs and associated 

research directions can be identified and considered 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 69

effectively and efficiently. Forests always will be in a 

state of transition and what we learned yesterday will 

be helpful but not totally applicable to the conditions 

today and in the future. We have no choice but to work 

toward sustaining our oak forests. We human beings 

have mediated global ecological changes and we have 

by no means grasped the magnitude of the unintended 

consequences of our actions. We are a significant part of 

the problem so we must diligently and deliberately be 

part of the solution. We should continue our efforts to 

sustain oak forests because:

1. The oaks have ecological “standing.” I borrow 

the word “standing” from the legal profession 

where it implies “a connection.” The oaks have 

been “connected” for millions of years. They 

have been displaced, replaced, and then returned 

over a notable length of geologic time. There 

is little question that they are of significant 

ecological importance in an evolutionary 

context.

2. There are at least 31 oak species in the Eastern 

United States and at least one oak species, (and 

often several species) is adapted to or will grow 

on virtually every forest site in the East.

3. The wood from most oak species is noted for its 

strength, durability, rot resistance, high energy 

value, and visual appeal that make it ideal for a 

host of important wood products. Think about 

a world without oak floors and fine furniture, 

railroad ties, bourbon and wine barrels, 

firewood, and pallets.

4. The oaks are among the longest living tree 

species in the East and, therefore, tend to 

stabilize forest communities that are subjected to 

outside forces—both natural and anthropogenic. 

The presence of oaks in forested communities 

tends to make those same communities more 

resilient.

5. The oaks provide essential habitat components 

for many wildlife species, including large and 

small mammals, birds, and reptiles. The volume 

and quality of food provided by acorns are 

essential for a host of species. The durability and 

size of oak cavity trees are seldom matched by 

associated tree species.

6. Oaks add color, shape, contrast and stability to 

virtually every landscape, in all four seasons and 

in both rural and urban settings.

7. The oak tree is a symbol of American forests. It 

has stature among tree species.

We should continue our efforts to sustain our oak forests 

and greatly expand our efforts to achieve that goal.

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Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 73

Vegetative and Forest 

Floor Responses to Fire
74	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Ecological and ecophysiological attributes and responses

to fire in eastern oak forests

Marc D. Abrams1

Abstract.—Prior to European settlement vast areas of the eastern U. S. deciduous forest were dominated 

by oak species. Evidence indicates that periodic understory fire was an important ecological factor in the 

historical development of oak forests. During European settlement of the late 19th and early 20th century, 

much of the Eastern United States was impacted by land clearing, extensive timber harvesting, severe fires, 

the chestnut blight, and then fire suppression and intensive deer browsing. These activities had the greatest 

negative impact on the once dominant white oak, while temporarily promoting the expansion of other oaks 

such as red oak and chestnut oak. More recently, recruitment of all the dominant upland oaks waned on all 

but the most xeric sites. Mixed-mesophytic and later successional hardwood species such as red maple, sugar 

maple, black birch, beech, black gum and black cherry are aggressively replacing oak. At Fort Indiantown 

Gap in southeast Pennsylvania, periodic burning over the last 50 years resulted in sites with lower tree 

density and a higher proportion of overstory oak species than unburned stands. Oak saplings averaged 

875 per ha in burned forests and 31 per ha in unburned forests. Red maple had overstory importance of 7 

percent and 24 percent in burned and unburned stands, respectively. The leaf litter of many oak replacement 

species (e.g., red maple, sugar maple, black birch, beech, black gum and black cherry) is less flammable 

and more rapidly mineralized than that of the upland oaks, reinforcing the lack of fire. The trend toward 

increases in non-oak tree species will continue in fire-suppressed forests, rendering them less combustible 

for forest managers who wish to restore natural fires regimes. Moreover, many of the oak replacement 

species are now growing too large, both above and below ground, to readily kill with understory fire. This 

situation greatly differs from the western United States, where fire suppression during the 20th century has 

made a variety of conifer-dominated forests more prone to stand-replacing fire. Thus, forest supervisors 

in the East who wish to use fire as a management tool in oak forests need to act sooner rather than later.

KEY WORDS: historical ecology, disturbance, succession, fire suppression, oak replacement

European settlement. These included extensive logging, 

land clearing, and catastrophic fire, followed by fire 

suppression and the introduction of exotic insects and 

diseases (Brose et al. 2001). All of these have led to 

unprecedented and rapid changes in forest composition 

and structure. This is particularly true for the Eastern 

United States, which has seen the extirpation of the once 

dominant chestnut (Castanea dentata) overstory from 

blight, loss of vast white pine (Pinus strobus) forests from 

logging followed by intense fires, a virtual cessation of 

oak regeneration from fire suppression and intensive 

deer browsing, and a rapid increase in native and exotic 

invasives (Keever 1953; Abrams 1992, 1998; Whitney 

1994). Some authors have characterized the landscape as 

undergoing a near complete transformation over the last 

350 years (Whitney 1994; Foster et al. 1998).

Oak was the dominant genus in the pre-European 

settlement forests throughout much of the Eastern 

United States (Abrams 1992; Whitney 1994). However, 

INTRODUCTION

The increased importance of oak (Quercus) during the 

Holocene epoch was associated with warmer and drier 

climate and elevated fire frequency after glacial retreat 

(reviewed in Abrams 2002). Significant levels of charcoal 

influx occurred almost routinely with oak pollen in lake 

and bog sediments throughout the Holocene (Delcourt 

and Delcourt 1997). As American Indian populations 

increased throughout the Eastern United States so did 

their use of fire, land clearing, and other agricultural 

activities (Whitney 1994). Thus, low to moderate levels 

of biotic and abiotic disturbance and climate change 

were an intrinsic part of the Holocene ecology, resulting 

in a dynamic equilibrium in regional forests.

The magnitude of anthropogenic disturbances in 

North American forests changed dramatically following 

1

Ferguson Building, School of Forest Resources, Pennsylvania 

State University, University Park, PA 16802, 814-865-4901, 

email: agl@psu.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 75

there has been little recruitment of oak during the 20th

century (Cho and Boerner 1991; Abrams et al. 1995), 

with the possible exception of the southwest portion 

of the eastern deciduous forest (parts of Missouri, 

Oklahoma, and Arkansas) that lacks many of the later 

successional, oak-replacement species (Abrams 1992). 

There is evidence of a dramatic decline in oak forests 

from presettlement to the present for much of the 

Eastern United States (Glitzenstein et al. 1990; Fralish 

et al. 1991; Whitney 1994; Abrams and Ruffner 1995). 

Anthropogenic impacts during the late 19th and early 

20th centuries included both the height and the end of 

the clearcutting era, catastrophic wildfires, the beginning 

of the fire suppression era, and chestnut blight. In 

this paper I synthesize studies of land-use history, 

witness trees (from early land surveys), dendroecology 

(tree-ring studies), and fire history to investigate the 

major ecological and environmental changes that have 

occurred in the eastern deciduous forest following 

European settlement. Due to the large increase in nonpyrogenic, mixed-mesophytic tree species, the window of 

opportunity to restore natural fire regimes in eastern oak 

forests may be closing in the foreseeable future.

OAK ABUNDANCE IN THE

PRESETTLEMENT FOREST

 Oak distribution in the pre-European settlement forests 

of New England differed dramatically from north to 

south. There is little or no oak in northern New England 

except along the Connecticut River Valley (Table 1; 

Burns and Honkala 1990; Cogbill et al. 2002). However, 

in southern New England and eastern New York, white 

oak was typically the first-rank species, with composition 

percentages ranging from about 17 to 36 percent 

(Table 1). Other dominant tree species in these forests 

included white pine, hickory (Carya), chestnut, and 

hemlock (Tsuga canadensis).

Oak species occurred in the southern and central regions 

of the Lake States (Burns and Honkala 1990). White oak 

represented 19 to 26 percent of certain presettlement 

forests in southern and central Michigan and Wisconsin, 

in some regions occurring with red pine (P. resinosa) and 

white pine, as part of bur oak (Q. macrocarpa) savannas, 

or with sugar maple (Acer saccharum) and red oak (Q. 

rubra) (Cottam 1949; Kilburn 1960; Nowacki et al. 1990)

The peak distribution for oak species in the 

presettlement forest clearly was in the oak-hickory, 

oak-pine, and former oak-chestnut regions of the MidAtlantic, central Appalachians and Piedmont, Midwest, 

and Central States (Table 1). In the Mid-Atlantic region, 

oak was the first- or second-rank species in 16 of 18 

studies reviewed here. In 12 of these examples, white oak 

was the dominant species, with a frequency of 21 to 49 

percent. White oak was second to black oak (Q. velutina; 

33 percent) with a frequency of 17 to 30 percent in the 

Piedmont of southeastern Pennsylvania (Mikan et al. 

1994; Black and Abrams 2001). In the Ridge and Valley 

of Pennsylvania, white oak was the first rank dominant 

on valley floors, but was a codominant behind pine 

species and chestnut oak on ridges (Nowacki and Abrams 

1992; Abrams and Ruffner 1995).

In the Midwest and central regions, white oak 

followed by black oak were the dominant species in 

six of eight examples (Table 1). However, fine till soils 

in northeastern Ohio and western New York were 

dominated by sugar maple and beech (Fagus grandifolia), 

with lesser amounts (5 to 14 percent) of white oak 

(Seischab 1990; Whitney 1994). White oak and black 

oak typically grew with hickory on drier and less fertile 

sites throughout the region.

There is much less information on pre-European forest 

composition for the South and Southeast (Table 1). The 

few existing studies suggest that oak species were not 

typically dominant but did achieve frequencies of 5 to 18 

percent in forests with Magnolia, beech, maple, pine and 

other oak species (Table 1). However, more numerous 

studies of 20th century forests and old-growth remnants 

farther north in the Piedmont and central and southern 

Appalachians suggest that oaks were a dominant in the 

original forest (Braun 1950, Peet and Christensen 1980, 

Monk et al. 1990; Barnes 1991).

OAK DECLINE FOLLOWING 

EUROPEAN SETTLEMENT

Significant changes in the composition of oak forests 

occurred in most regions from presettlement the to 

present. In 18 of 26 examples reviewed here, white oak 

experienced a decline in frequency of 10 percent or more 

(Table 2). Six examples reported no significant change, 
76	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Table 1.—Percent composition of witness tree species in pre-European settlement forests in the Eastern United States

Region and location Presettlement forest composition Reference

Northeast

northern VT, NH beech(32), spruce (14), maple (12), hemlock (12), oak (5) Cogbill et al. 2002

western NY beech (32), sugar maple (18), basswood (12), white oak (11) Seischab 1990

eastern NY white oak (36), black oak (15), hickory (10), elm (6) Glitzenstein et al. 1990

central MA white oak (27), black oak (26), pine (18), hickory (9) Whitney and Davis 1986

central MA white oak (20), pine (20), hemlock (10), chestnut (8) Foster et al. 1998

CT and RI white oak (33), hickory (10), chestnut (9), Cogbill et al. 2002

MA white oak (25), pine (16), maple (6), hemlock (6) Cogbill et al. 2002

eastern NY white oak (17), beech (16), hemlock (10), pine (9) Cogbill et al. 2002

Lake States

central MI jack pine (20), red pine (19), white pine (11), white oak (2) Whitney 1994

central MI red pine (40), white oak (19), white pine (15), aspen (12) Kilburn 1960

southern WI bur oak (60), white oak (26), black oak (13) Cottam 1949

central WI sugar maple (37), white oak (25), red oak (16), elm (12) Curtis 1959

central WI pine (28), aspen (17), larch (12), white oak (10) Nowacki et al. 1990 

Mid-Atlantic

northern NJ white oak (34), black oak (18), hickory (15), red oak (9) Russell 1981 

northern NJ white oak (31), hickory (25), black oak (19), chestnut (12) Ehrenfeld 1982

northwest PA white oak (21), beech (13), maple (17), black oak (6) Whitney and Decant 2001

southeast PA black oak (33), white oak (17), chestnut (15), hickory (15) Mikan et al. 1994

southeast PA black oak (33), white oak (30), hickory (28) Black and Abrams 2001

southwest PA white oak (40), black oak (9), hickory (9), dogwood (8) Abrams and Downs 1990

central PA

 Allegheny Mts

 plateaus white oak (26), chestnut (19), pine (19), maple (10) Abrams and Ruffner 1995

 stream valleys hemlock (24), maple (21), white pine (15), birch (15) Abrams and Ruffner 1995

 Ridge and Valley

 ridges chestnut oak (14), white oak (12), pine (19), chestnut (11) Abrams and Ruffner 1995

 valleys white oak (30), pine (25), hickory (17), black oak (10) Abrams and Ruffner 1995

 ridges pine (27), chestnut oak (18), white oak (11), chestnut (13) Nowacki and Abrams 1992

 valley white oak (41), white pine (12), hickory (12), black oak (9) Nowacki and Abrams 1992

eastern WV

 ridges white oak (35), chestnut (15), chestnut oak (13), black oak (12) Abrams and McCay 1996

 valleys white oak (23), maple (22), pine(15), basswood (10) Abrams and McCay 1996

southern WV white oak (24), chestnut (12), hickory (9), chestnut oak (6) Abrams et al. 1995

northern VA white oak (49), red oak (26), hickory (7) Orwig and Abrams 1994

southwest VA red oak (25), white oak (18), chestnut (9) McCormick and Platt 1980

western Virginia white oak (26), pine (13), chestnut oak (9), hickory (9) Stephenson et al. 1992

Mid-west and central region

central MO white oak (32), black oak (11), sugar maple (9), elm (8), Wuenscher and Valiunas 1967

eastern IL white oak (27), black oak (18), hickory (6), elm (10) Rodgers and Anderson 1979

southern IL

 south slopes white oak (81) Fralish et al. 1991

 ridge tops white oak (45), black oak (33), Fralish et al. 1991

northeast OH

 fine till beech (36), sugar maple (17), white oak (14) Whitney 1994

 coarse till white oak (37), hickory (13), black oak (6) Whitney 1994

north-central OH hickory (34), white oak (30), bur oak (11), black oak (11) Whitney 1994

southeast OH white oak (40), hickory (14), black oak (12), beech (8) Dyer 2001

South and southeast

north FL magnolia (21), beech (14), maple (7), white oak (5) Delcourt and Delcourt 1977

central GA pine (27), black and red oak (21), post oak (18), white oak (7) Cowell 1995

southeast TX pine (25), white oak (18), pin oak (10), red oak (9) Schafale and Harcombe 1983

eastern AL white oak (13), beech (9), pine (9), maple (5) Black et al. 2002
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 77

whereas two cases actually showed an increase or more 

than 10 percent for the species. The latter examples are 

rather special cases that involved the conversion of bur 

oak savannas to closed oak forests in southern Wisconsin 

following Euro-American settlement and fire suppression 

(Cottam 1949; Whitney 1994).

The magnitude of decline in the once dominant 

white oak has been dramatic. For example, in central 

Massachusetts oak (mainly white oak) decreased in 

frequency by more than 20 percent in the Connecticut 

Valley and Eastern Lowlands (Table 2; Foster et al. 1998). 

In the Hudson Valley of eastern New York, white oak 

declined mroe than 30 percent (Glitzenstein et al. 1990). 

Similar declines were noted for white oak in northwest 

and southeastern Pennsylvania, eastern West Virginia, 

and northern Virginia (Orwig and Abrams 1994; Abrams 

and McCay 1996; Black and Abrams 2001; Whitney and 

Table 2.—Percent frequency of present-day forest composition in the Eastern United States (examples chosen based on 

availability of corresponding pre-European settlement witness tree data)

Region and location Present day Reference

Northeast

MA 

 Connecticut Valley maple (30), oak (22), hemlock(15), pine (11), birch (11) Foster et al. 1998

 Pellham Hills maple (27), oak (21), hemlock(15), birch (15), pine (11) Foster et al. 1998

 Central Uplands maple (24), oak (23), pine(16), birch (12), hemlock (12) Foster et al. 1998

 Eastern Lowlands oak (35), maple (23), pine(21), birch (8) Foster et al. 1998

central MA white pine (23), black oak (21), red oak (19), white oak (9) Whitney and Davis 1986

eastern NY maple (30), chestnut oak (14), red oak (10), pine (9), white oak (4) Glitzenstein et al. 1990

Mid-Atlantic

northwest PA red maple (22), black cherry (14), hemlock (7), white oak (3) Whitney and Decant 2001

southeast PA chestnut oak (26), red maple (18), black oak (15), white oak (4) Black and Abrams 2001

southeast PA box elder (23), red maple (19), ash(8), elm (7), white oak (1) Kuhn (unpublished data)

central PA

 Allegheny Mts. red maple (35), white oak (19), red oak (11), chestnut oak (9) Abrams and Ruffner 1995

 Ridge and Valley chestnut oak (28), red maple (14), red oak (14), white oak (13) Abrams and Ruffner 1995

 valleys white oak (43), red maple (15), black cherry (10), pine (7) Nowacki and Abrams 1992

 ridges chestnut oak (43), red oak (19), red maple (14), white oak (1) Nowacki and Abrams 1992

southwest PA red maple (30), beech (23), tulip poplar (17), white oak (5) Abrams and Downs 1990

eastern WV chestnut oak (15), red oak (14), red maple (12), white oak (9) Abrams and McCay 1996

northern VA white oak (30), hickory (13), poplar (13), dogwood (11) Orwig and Abrams 1994

southwest VA hickory (14), red oak (12), chestnut oak (8), white oak (5) McCormick and Platt 1980

Midwest and Lake States

southeast OH white oak (15), black oak (14), tulip poplar (11), hickory (8) Dyer 2001

northeast OH beech (11), white oak (11), hickory (9), black cherry (8), red maple (6) Whitney 1994

southern WI sugar maple (28), elm (14), basswood (11), white ask (10), white oak (5) Whitney 1994

southern WI (savanna)white oak (34), hickory (30), black oak (24), black cherry (12) Whitney 1994

southern WI (savanna)white oak (54), black oak (25), black cherry (17) Cottam 1949

central Wisconsin red oak (46), white oak (19), red maple (16) Nowacki et al. 1990

southern IL

 south slope white oak (30), black oak (22), post oak (18), hickory (13) Fralish et al. 1991

 ridge top white oak (53), black oak (17), hickory (14), post oak (7) Fralish et al. 1991

 north slope white oak (21), red oak ( 22), sugar maple (13), black oak (13) Fralish et al. 1991
78	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Decant 2001). However, the largest decline in white oak 

(from 81 percent to 30 percent) was reported on south 

slopes in southern Illinois (Fralish et al. 1991).

Currently, oak is the first rank tree species in only 14 of 

the 26 examples reviewed here compared to 24 of these 

examples at the time of European settlement (Tables 

1 - 2). There is a greater tendency for present-day oak 

dominance in the Midwest and Lake States, where 

they increased in former bur oak savannas and logged 

and burned-over pine forests (Cottam 1949; Whitney 

1994; Nowacki et al. 1990), or in the former prairie 

peninsula outside the range of red maple (Fralish et al. 

1991). Apart from these exceptions, white oak generally 

experienced a significant decline in frequency even when 

it maintained the dominant ranking in modern forests 

(Fralish et al. 1991; Orwig and Abrams 1994; Foster et 

al. 1998; Dyer 2001).

Large increases in red oak and chestnut oak also occurred 

from presettlement to the present (Tables 1 - 2). Red oak 

increased from 7 to 19 percent in central Massachusetts 

and from 2 to 22 percent on north-facing slopes in 

southern Illinois (Fralish et al. 1991; Whitney and 

Davis 1986). Red oak has importance values of 37 to 51

percent in present forests of north-central Wisconsin, 

where it formerly represented less than 1 percent of the 

original northern hardwood-conifer forest (Nowacki 

et al. 1990). Increases in red oak ranging from 9 to 

19 percent and in chestnut oak from 7 to 25 percent 

occurred in the Allegheny Mountains and Ridge and 

Valley of Pennsylvania and West Virginia (Nowacki and 

Abrams 1992; Abrams and Ruffner 1995; Abrams and 

McCay 1996). In western Virginia, red oak increased 

from 11 to 57 percent (from 1932 to 1982), while red 

maple increased from 1 to 11 percent (Stephenson et al. 

1992). Chestnut oak increased in frequency from less 

than 1 percent to 26 percent in the Piedmont lowlands 

of southeastern Pennsylvania (Black and Abrams 2001). 

Red oak and chestnut oak apparently benefited from the 

death of overstory chestnut (on ridges), selective logging 

of white oak, and the intensive logging and burning of 

both high- and low-elevation forests.

The witness tree studies reviewed here indicate that 

the once dominant white oak grew on a wider range of 

sites and in greater numbers than any other eastern oak 

species (Braun 1950; Peet and Christensen 1980; Monk 

et al. 1990; Barnes 1991). Red oak appears frequently 

in the witness tree record but only occasionally had a 

frequency exceeding 5 percent. It is likely that there were 

small populations of red oak across the eastern forest on 

most landforms that provided adequate nutrients and 

some protection from fire and drought.

After the clearcut era of the late 1800s and the 

chestnut blight of the early 1900s, the fast-growing and 

opportunistic red oak expanded dramatically from its 

sheltered areas and grew over vast areas of the eastern 

forest previously dominated by chestnut, pine, and white 

oak (Keever 1953; Crow 1988; Nowacki et al. 1990; 

Barnes 1991; Stephenson et al. 1992). It appears that red 

oak flourished in response to large-scale anthropogenic 

disturbances that were much less common in the preEuropean settlement forest. Moreover, the invasion of 

red oak into relatively undisturbed old-growth white 

oak forests probably was facilitated by its increase in 

surrounding forests that were more highly disturbed 

(Abrams et al. 1995; Abrams and Copenheaver 1999).

The post-settlement expansion of chestnut oak can be 

best explained by the loss of overstory chestnut, pitch 

pine (Pinus rigida) and white oak, and it being tolerant 

of severe fires that occurred during and after the major 

clearcut era in the late 1800s (Keever 1953; McCormick 

and Platt 1980; Abrams and Ruffner 1995; Abrams and 

McCay 1996). Despite often growing on ridge sites and 

being a low quality timber species, chestnut oak did not 

escape extensive cutting during the 19th century. It was an 

important fuelwood for domestic uses and the charcoal 

iron industry, and a source of tannin for the tanbark 

industry (Stephenson et al. 1992; Whitney 1994). On 

sites in which chestnut oak and white oak codominated 

in the original forest, chestnut oak was the apparent 

victor following the catastrophic disturbances. However, 

many chestnut oak forests now are being invaded by red 

maple (Table 2).

RISE IN RED MAPLE IN EASTERN OAK

FORESTS

By far, the largest increases in species frequency on 

present-day upland oaks sites are from red maple (Tables 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 79

1 - 2). Red maple now represents the first or 

second rank dominant in 12 of the 17 examples 

from southern New England, eastern New York, 

and the mid-Atlantic region. This is even more 

impressive when you consider that little red 

maple was recorded in the presettlement forests 

of these areas (Table 1). Moreover, many oldgrowth oak forests now have abundant young red 

maple as a dominant tree (Abrams and Downs 

1990; Mikan et al 1994; Shumway et al. 2001; 

Abrams et al. 1995). The dramatic rise in red 

maple in oak forests during the 20th century has 

been attributed to the extensive logging of oak in 

the late 19th and early 20th century, the chestnut 

blight, and the suppression of understory burning 

(Abrams 1992, 1998; Nowacki and Abrams 

1992; Stephenson et al. 1992; Mikan et al. 

1994). In the prairie regions of Illinois, Iowa, and 

Missouri, outside the range of red maple, sugar 

maple now is the dominant later successional, oak 

replacement species on mesic, nutrient-rich sites 

(Pallardy et al. 1988; Fralish et al. 1991).

The situation for red maple is somewhat analogous 

to the rise in red oak during the 20th century in the 

Eastern United States (Abrams 1998). Before European 

settlement, red maple generally was limited to swamps 

and other areas sheltered from fire. Red maple is much 

more sensitive to fire than red oak, and would have 

been less common on uplands. After 1900, red maple 

quickly expanded out of the protected areas and began to 

dominate most forest understories throughout its range. 

The selective logging of the highly prized white oak gave 

a clear opportunity to less common and/or less desirable 

species, e.g., red oak, red maple, black birch, and black 

cherry (Whitney 1994; Whitney and Decant 2001).

A CASE STUDY OF FIRE HISTORY 

AND OAK RECRUITMENT IN AN 

OLD-GROWTH OAK FOREST

Fire history and dendroecology (based on tree ring 

analysis) were investigated for two stands in an oldgrowth, mixed-oak stands in western Maryland 

(Shumway et al. 2001). I believe the ecological history 

and dynamics of these stands are representative of oak 

forests throughout much of the eastern forest. One 

stand (Coleman Hollow) is dominated by red maple 

(24 percent), chestnut oak (23 percent), white oak (20 

percent), red oak (14 percent), and black oak (9 percent), 

whereas the South Savage stand is dominated by 

chestnut oak (20 percent), black birch (Betula lenta; 18 

percent), red oak (17 percent), red maple (17 percent), 

black oak (11 percent), and white oak (6 percent). A 

greater abundance of rock outcroppings at South Savage 

may have allowed for more black birch and less white 

oak in the stand. The presettlement forest on Savage 

Mountain contained white oak (27 percent), hickory 

(18 percent), black oak (12 percent), chestnut oak (11

percent), chestnut (10 percent), and red oak (5 percent).

Basal cross sections were obtained from a partial timber 

cut in 1986, which provided evidence of 42 fires from 

1615 to 1958. Fires occurred on average every 8 years 

during the presettlement (1600 to 1780) and early postsettlement (1780 to 1900). These included seven major 

fires year in which at least 25 percent of the sample trees 

were scarred in a given year (Fig. 1). No major fire years 

occurred after 1900 and no fires were recorded after 

1960. The South Savage stand had a larger component 

Figure 1.—Age-diameter relationships for all cored trees and the major 

fire years (> 25% of sampled trees scarred; indicated by triangles along 

x-axis) in two old-growth forests on Savage Mountain in western Maryland 

(adapted from Shumway et al. 2001). WO = white oak; CO = chestnut 

oak, BO = black oak, BB = black birch, RM = red maple, PH = pignut 

hickory, OT = other tree species.
80	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

of older trees, including a 409-year-old white oak, and 

exhibited continuous recruitment of oaks from the late 

1500s until 1900. Interestingly, white oak and chestnut 

oak dominated recruitment from 1650 to 1800, whereas 

red oak and black oak dominated recruitment from 1800 

to 1900. The lack of red oak and black oak recruitment 

prior to 1800 may be due to their relatively short 

longevity at the site. However, it is difficult to explain 

the large reduction in white oak and chestnut oak 

recruitment after 1800, though they might have been 

outcompeted by the other oaks. After 1900, red oak was 

the only oak species to recruit in significant numbers. 

This was associated with the loss of overstory chestnut 

from the blight.

Coleman Hollow differed from South Savage in species 

composition and the fact that it contained only two very 

old oaks—white oaks 290 and 320 years old (Fig. 1). 

Moreover, the abundant oak recruitment during the 

19th century included large amounts of chestnut oak 

and white oak not seen on Savage South. From 1900 

to 1950, recruitment of chestnut oak and red oak was 

joined by large numbers of red maple and black birch. 

There was little recruitment of white oak in either stand 

after 1900.

The results of this study indicate that periodic fires 

burned through the forest understories between 1600 

and 1900, and that some degree of burning continued 

until 1960 because of its remote location. The fire 

rotation at Savage Mountain is consistent with mean 

fire intervals of 4 to 20 years in other oak forests in the 

Eastern and Central United States (Guyette and Dey 

1995; Sutherland 1997; Schuler and McClain 2003). 

The long history of periodic burning at the study site 

was associated with continuous oak recruitment. A 

similar result was reported for a fire history and red 

oak recruitment study in West Virginia (Schuler and 

McClain 2003). Fires at Savage Mountain likely played 

an important role in oak ecology, such as preparing a 

thin litter seedbed, increasing sunlight to the forest floor, 

and suppressing red maple and black birch. Indeed, red 

maple and black birch were absent from the witness tree 

record and among the older trees in the forests, even 

though they can live for more than 200 years. Large 

amounts of red oak, chestnut oak, red maple, and birch 

recruitment were associated with the chestnut blight 

period from 1910 to 1950. A reduction and eventual 

cessation of fire further facilitated red maple and 

black birch invasion in the forest while retarding the 

recruitment of all oak species.

IMPACTS OF RECENT, RECURRING 

FIRE ON OAK FOREST COMPOSITION 

AND RRECRUITMENT

The National Guard Training Center at Fort Indiantown 

Gap near Harrisburg, Pennsylvania, has experienced 

frequent fires since the 1950’s on the ridges and since 

the 1980’s in the valleys as a result of military training 

exercises. This represented an unique opportunity to 

investigate the role of recent and repeated fire in oak 

forests in the Eastern United States. We investigated four 

frequently burned and two unburned sites replicated in 

Ridge and Valley ecosystems (Signell et al. 2006).

The cross sections collected from stand VB1 indicate 

that it initiated following a fire in 1941; a second fire 

occurred in 1953 (Fig. 2). Most cross sections had 

pith dates of 1953 but several individuals initiated in 

1942 and one of these had a 1953 fire scar. VB1 has 

also experienced major fires in 1984, 1986, 1990 and 

1992, with mean fire return interval (FRI) of 2.7 years 

for the period. The diameter distribution of VB1 is bell 

shaped rather than positively skewed as in VU (Fig. 3). 

Oak dominated every size class and shade-tolerant/fire 

sensitive species were uncommon. Total oak importance 

value exceeded 94 percent in this stand. Cross sections 

from stand VB2 showed recruitment dates from the 

early 1940s. Like VB1, this stand experienced a fire-free 

period following stand initiation. No fire scars were 

found until the early 1980s, after which at least three 

fires occurred, in 1982, 1985 and 1990, resulting in a 

mean FRI of 4 years. VB2 has a bell-shaped diameter 

distribution similar to that for VB1, and oak dominated 

all size classes (Fig. 3). However, VB2 has more shade 

tolerant-species present in the smaller diameter classes, 

including red maple and blackgum, than were observed 

in stand VB1. Total oak importance value in stand VB2

was 81.1 percent.

Stands RU and RB1 were part of an uninterrupted 

forest stand in 1952. Tree-ring data show that the 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 81

oldest trees in both stands originated around 1845 

(Fig. 2). No fire scars were found in any of the cross 

sections obtained from these stands until 1952, when 

both stands experienced a fire severe enough to scar 

many of the approximately 100-year-old chestnut oaks 

present at the time. Between 1952 and 1956, a road was 

constructed between the two stands. Following the road 

construction, RB1 experienced fires in 1979, 1983 and 

1991, while RU has been free of fire (Fig.2).

RU had high stem density, mostly from chestnut oak, 

red maple, and black birch (Fig. 4), but few individuals 

more than 30 cm in diameter at breast height, and they 

were mostly chestnut oak. Nonetheless, these large trees 

contributed greatly to the high basal area in this stand. 

Red maple and black birch dominated the smaller size 

classes (Fig. 4). RB1 had lower stem density than stand 

RU. Of the major species, only scarlet oak had more 

stems in stand RB1 versus RU. Most black birch and 

red maple were concentrated in rocky patches. Chestnut 

oak had the highest importance in this stand; overall oak 

importance was 63.7 percent. Stand RB2 had several 

large chestnut oaks and red oaks that were clustered on 

a rocky boulder field. Fire occurred in 1890 and 1900, 

soon after stand establishment (Fig. 2). No fire scars 

were recorded between 1900 and 1952. Many of the 

smaller trees recruited following a fire in 1953. RB2 had 

additional fires in 1986, 1991, 1996, and 2000 (Fig. 2). 

Figure 2.—Fire scar records obtained from tree basal cross sections at 

Fort Indiantown Gap, Pennsylvania. Horizontal lines represent the visible 

growth rings for each tree, and vertical bars represent fire scars (adapted 

from Signell et al. 2005).
8	 Fire 2 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

percent relative importance value) than unburned stands 

(47 to 49 percent importance). Oak saplings averaged 

875 per ha in burned forests and 31 per ha in unburned 

forests. Red maple, the most aggressive oak replacement 

species in the Eastern United States, had overstory 

importance of 7 percent and 24 percent in burned and 

unburned stands, respectively. Oak saplings ranged 

from 824 to 1,545 per ha in three of the four burned 

Figure 3.—Diameter distributions for unburned (VU) and burned valley stands 

(VB1, VB2) at Fort Indiantown Gap, Pennsylvania. Quve = Quercus velutina, Qual 

= Quercus alba, Quru = Quercus rubra, Qupr = Quercus prinus, Quco = Quercus 

coccinnea, Nysy = Nyssa sylvatica, Bele = Betula lenta, Acru = Acer rubrum

(adapted from Signell et al. 2005).

RB2 has high tree density, with oak species dominating 

all diameter classes (Fig. 4). There were substantial 

numbers of red maple and black birch, primarily on 

rocky patches. Oak importance was 69.2 percent in this 

stand.

Burned sites generally had lower tree density and a 

higher proportion of overstory oak species (64 to 92
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 83

stands and 0 to 62 per ha in the unburned stands. Oak 

sapling density was only 62 per ha in one recently (2002) 

burned ridge stand that had the highest tree density. 

There were no red maple saplings in three of the four 

burned stands. Oak saplings were most abundant when 

overstory density was less than 400 trees per ha and 

understory tree density was less than 200 trees per ha. 

When overstory or understory tree density exceeded 

400 and 200 trees/ha respectively, oak regeneration was 

virtually absent. The results of this study suggest that 

periodic fire often reduces overstory and understory 

stand density and promotes successful regeneration of 

relatively shade intolerant oak species in the Eastern 

United States. However, high tree density in forests 

will retard the development of oak understories and 

subsequent recruitment even if periodic burning occurs.

Figure 4.—Diameter distributions for unburned (RU) and burned ridge stands 

(RB1, RB2) at Fort Indiantown Gap, Pennsylvania. Quve = Quercus velutina, 

Qual = Quercus alba, Quru = Quercus rubra, Qupr = Quercus prinus, Quco = 

Quercus coccinnea, Nysy = Nyssa sylvatica, Bele = Betula lenta, Acru = Acer 

rubrum (adapted from Signell et al. 2005).
84	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

ECOPHYSIOLOGICAL ATTRIBUTES OF 

UPLAND OAK SPECIES

Upland oaks are prevalent across a broad array of sites, 

including drought-prone areas, have evolved with 

periodic understory burning and are transitional to 

later successional species in the absence of fire on most 

sites. Thus, upland oaks presumably possess a suite of 

ecophysiological adaptations for drought and fire but not 

for competing in a closed forest understory dominated 

by more shade-tolerant species. In this section I 

summarize the ecophysiological attributes of upland oaks 

as they relate to its success in the presettlement forest 

and subsequent decline following European settlement 

(Table 3).

Fire Adaptations

In an early opinion survey of the fire resistance for 22

northeastern tree species, oaks (chestnut oak, black oak, 

white oak, and scarlet oak) were rated in four of the top 

six positions (Starker 1934). A ranking of increasing bark 

thickness and fire resistance reported that bur oak > black 

oak > white oak > red oak (Lorimer 1985). White oaks 

(subgenus Leucobalanus), including bur oak, chestnut 

oak, and white oak, can produce tyloses, eccentric 

outgrowths of cell walls in response to wounding, 

which allow for effective compartmentalization of fire 

scar injuries. In this respect, white oaks should be more 

resistant to fire than red oaks (Table 3). Fire also may 

be beneficial to oaks relative to other hardwood species 

because of their deep and extensive rooting, vigorous 

sprouting ability, and increased germination and survival 

on fire-created seedbeds (Table 3).

Drought Adaptations

Oaks are among the most deeply rooted tree species 

in the Eastern United States, which allows them to 

maintain relatively high predawn shoot water potential 

from superior overnight rehydration (Table 3). Oak 

leaves typically are more xerophytic, having greater 

Table 3.—Adaptations to fire, drought and understory conditions in upland oaks relative to non-oak hardwood 

species (adapted from Abrams 1996)

Fire Drought Understory

Thick bark Deep rooting Low-moderate shade tolerance

Rotting resistance after

 scarring; tyloses in white oaks

High predawn leaf water potential Large seed and cotyledons

Deep and extensive rooting Thick leaves Large initial seedling and roots

Vigorous sprouting in young trees High leaf mass per area Relatively high net photosynthesis

Improved fire-created seedbed High stomatal density Relatively low respiration

Increased germination High leaf nitrogen Low light compensation point

Increased seedling survival High net photosynthesis Medium light saturation values

Lowered seed and seedling predation High leaf conductance Slow seedling shoot growth

High drought threshold for

 stomatal closure

High c-based phenolics

High osmotic adjustment Shoot dieback

High wilting threshold Intense seed and seedling predation

Low nonstomatal inhibition of 

 photosynthesis

Often overtopped by competing species

Leaves not drought deciduous Responsive to canopy gaps

Increase in mixed-mesophytic hardwoods

 decreasing litter layer flammability 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 85

thickness, mass per area and stomatal density, and higher 

nitrogen content than leaves of non-oak species. These 

factors contribute to the relatively high photosynthesis 

and transpiration in white oak and other oak species. 

Oaks need to develop the necessary tissue water relations 

to support high level of gas exchange, and they often 

exhibit low osmotic potentials, low relative water content 

at zero turgor, and low water potential threshold for 

stomatal closure than non-oak species (reviewed in 

Abrams 1992).

Adaptations to Forest Understory 

Conditions

Most upland oaks are considered to have intermediate 

shade tolerance (Table 3; Burns and Honkala 1990), 

though Crow (1988) considered red oak to have low 

shade tolerance. Oaks produce a large acorn that allow 

for high initial growth, though white oaks typically 

exhibit low shoot growth after the first year in forest 

understories (Cho and Boerner 1991). Sapling density 

is low for most oaks, especially for white oak. This 

indicates that there is a severe “bottleneck” between the 

oak seedling and sapling stages (Crow 1988; Nowacki et 

al. 1990).

The physiological mechanisms for slow growth in oak 

seedlings are complex. Photosynthetic rates of oak in 

shaded conditions are often higher than in non-oak 

species, while oak respiration rates are low to moderate 

(Table 3). Oak seedlings often produce large root systems 

but experience recurring partial or complete shoot 

dieback (Crow 1988; Abrams 1992, 1996). Allocating 

carbon in this manner has its limitation, particularly for 

the once dominant white oak. White oak is reported to 

have slower height growth than chestnut, red oak, and 

black oak (Whitney 1994). Following a 1985 tornado, 

white oak seedling/sprouts had among the lowest rates of 

height growth for seven species (including black oak, red 

oak, and black cherry; Kauffman 2002).

Upland oaks are well adapted to dealing with a range 

of moisture conditions, and their moderate shade 

tolerance is conducive to gap-phase regeneration in 

fire-maintained forests. However, upland oaks grow 

slowly in deeply shaded forest understories dominated by 

later successional species and in heavily disturbed areas, 

where they are overtopped by competitors. Intensive 

deer browsing, despite the production of carbon-based 

defense chemicals, only exacerbates the situation (Table 

3). Moreover, the increase in later successional, mixedmesophytic hardwoods is reducing the flammability of 

the litter layer, which likely will result in less frequent 

and intense surface fires in the future. These ecological 

stressors represent a serious threat to the survival of 

upland oaks.

CONCLUSION

Paleoecological and dendroecological evidence suggest 

that the process of fire and oak recruitment in upland 

forests went on for many hundreds and thousands of 

years. This cycle was broken during the 19th and early 

20th century, leading to a dramatic decline in the once 

super-dominant white oak followed by declines in 

other oak species. It is doubtful that oaks on mesic sites 

represent a true self-perpetuating climax in the absence 

of fire (Lorimer 1985; Abrams 1992; Schuler and 

McClain 2003). Thus, the broad ecological distribution 

of oaks in the Eastern United States can be attributed 

directly (probably in large part) to extensive understory 

burning in the pre-European and early settlement forests. 

If not for these fires, most oak sites would have been 

dominated by red maple, sugar maple, birch, beech, and 

black gum, a trend that now is apparent throughout 

most of their range.

There has been a nearly complete cessation of oak 

recruitment over the last 50 to 100 years in all but the 

most xeric and nutrient poor sites. There now is strong 

competitive pressure on oak regeneration from a number 

of later successional and gap opportunistic trees. The 

conversion of flammable oak litter, with high lignin 

content, in forest understories to less combustible and 

more rapidly decomposed litter of mixed-mesophytic 

and later successional tree species (Melillo et al. 

1982; Lorimer 1985; Washburn and Arthur 2003) is 

rendering eastern oak forests less prone to burning. 

By contrast, fire suppression in the Western United 

States and resultant increases in live and dead fuel, 

stand density, and changes in species composition have 

made many conifer-dominated forests more prone to 
86	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

fire (Biswell 1967; Parsons 1976; Brown et al. 2000). 

The declining combustibility of eastern forests may be 

further exacerbated by intensive deer browsing, which 

has reduced the leaf litter from most woody species, 

especially oaks. Thus, forest managers wishing to restore 

historical burning regimes to eastern forests in the hope 

of encouraging more oak regeneration while reducing 

native invasive tree species should act sooner rather than 

later as the window of opportunity may be closing.

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90	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Research Efforts on Fuels, Fuel Models, and Fire Behavior

in Eastern Hardwood Forests

Thomas A. Waldrop1†, Lucy Brudnak1, Ross J. Phillips1, and Patrick H. Brose2

Abstract.—Although fire was historically important to most eastern hardwood systems, its 

reintroduction by prescribed burning programs has been slow. As a result, less information 

is available on these systems to fire managers. Recent research and nationwide programs are 

beginning to produce usable products to predict fuel accumulation and fire behavior. We introduce 

some of those tools and examine results from regional studies of hardwood fuel distribution.

INTRODUCTION

A broad array of natural disturbances has contributed to 

the development of eastern hardwood ecosystems. The 

role played by natural and anthropogenic fire has not 

been appreciated until recent years. There is considerable 

knowledge on the fire-dependent pine ecosystems of 

the Western United States and the Southeastern Coastal 

Plain. However the role of fire is not well recognized for 

the Central Hardwood region, Southeastern Piedmont, 

and Southern Appalachian Mountains. In these areas, 

prescribed burning programs and supporting research has 

lagged behind that of the West and Southeastern Coastal 

Plain because of the perceived damage to hardwoods, 

difficulty of controlling high-intensity fires on slopes, 

and the potential for soil/site damage (Van Lear and 

Waldrop 1989). Consequently, basic fire-planning 

information such as fuel models and fire behavior models 

are missing or poorly calibrated.

Although fire was never entirely missing from the eastern 

hardwood region, prescribed burning only began to 

be used in hardwood stands during the 1980’s for site 

preparation (Phillips and Abercrombie 1987) and in the 

1990’s for restoration of individual species (Waldrop 

and Brose 1999) or entire ecosystems (Sutherland and 

Hutchinson 2003). Fire managers use a number of tools 

to describe fuels and predict fire behavior. Many of these 

tools were developed for western systems and have not 

been tested for eastern hardwoods. The purpose of this 

1

USDA Forest Service, Southern Research Station, 233 

Lehotsky Hall, Clemson, SC 29634-0331; 2USDA Forest 

Service, Northern Research Station, PO BOX 267, Irvine 

PA 16329; †Corresponding author, 864-656-5054, email: 

twaldrop@fs.fed.us.

paper is to examine several existing models of fuels and 

fire behavior, describe several ongoing projects that will 

develop new models, and describe fuel characteristics 

specific to the eastern hardwood region.

National Projects with Hardwood 

Fuel Components

Models have been developed for application in numerous 

ecosystems throughout the United States. Probably the 

most well known and used in eastern hardwood systems 

are BEHAVE (Burgan and Rothermel 1984), FARSITE 

(Finney 1998), and FOFEM (Reinhardt and others 

1997). Newer projects include LANDFIRE (http://www.

landfire.gov), a photo series for major natural fuel types 

of the United States (http://www.fs.fed.us/pnw/fera/), 

and the USDA Forest Service Forest Inventory and 

Analysis (FIA) National Assessment of Fuel Loadings 

(Woodall 2003). We present a brief description of 

each that is intended as an introduction rather than 

an exhaustive evaluation of all available fuel and fire 

behavior models.

BEHAVE and BEHAVEPlus

The BEHAVE fire prediction system was developed 

by Forest Service researchers to predict fire spread 

and intensity (Burgan and Rothermel 1984; Andrews 

1986; Andrews and Chase 1989; Andrews and 

Bradshaw 1990). Three later versions of this model 

(BEHAVEPlus v.1, v.2, and v.3) were developed to 

allow use in a Windows®-based environment and to 

incorporate updates in fire and modeling technology. 

This series of programs uses 13 fuel models that describe 

common fire-prone fuel types like grasslands, chaparral, 

coniferous forests, and logging slash (Anderson 1982). 

A set of 40 additional models, described by Scott and 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 91

Burgan (2005), was added in the most recent version 

of BehavePlus (3.0.1) that allow both static and 

dynamic simulations (that is, simulations that involve 

seasonally changing fuelbeds). The most commonly 

used fuel models for eastern hardwood systems are 

models 6 (hardwood slash), 8 (closed timber litter), 

and 9 (hardwood litter) (Brose 1997). The system also 

allows input of local fuel measurements by developing 

a customized fuel model. These are used for areas 

where fuel conditions are not adequately described by 

the standardized fuel models (Burgan and Rothermel 

1984). A description of each version of BEHAVE and 

BEHAVEPlus is found at http://fire.org. Free program 

downloads are available at this site.

Published tests of BEHAVE in eastern hardwood systems 

are rare but have been presented by Brose (1997) for 

oak-dominated (Quercus sp.) shelterwood stands in 

central Virginia and by Grabner and others (1997) for 

oak savannas in Missouri. Both studies used the original 

BEHAVE model. No published tests of the 40 new fuel 

models are available for eastern hardwood systems. Brose 

(1997) compared actual flame length (FL) and rate of 

spread (ROS) measurements from winter, spring, and 

summer burns to values predicted by BEHAVE using 

fuel models 6, 8, 9, and a customized fuel model. Of 

the four, the customized fuel model most accurately 

predicted both flame length and rate of spread. Model 

6 overestimated these measures of fire behavior while 

models 8 and 9 consistently underestimated them. 

The custom fuel model provided accurate predictions 

with r2 values of 0.88 for flame length and 0.96 for rate 

of spread. Grabner and others (1997) also compared 

observed and predicted values for FL and ROS. 

They tested BEHAVE fuel models 1 (short grass), 2

(timber and grass), 3 (tall grass), 9 (hardwood litter), 

and a customized fuel model. In this comparison, the 

customized fuel model and fuel model 2 provided 

reasonable estimates of ROS but all models poorly 

estimated FL. In both studies, the authors concluded 

that BEHAVE was a useful tool for eastern hardwood 

systems, especially when site-specific fuel measurements 

can be used as input for a customized fuel model. When 

those types of measurements are impossible, BEHAVE 

fuel models can be substituted with a reasonable degree 

of confidence.

A limitation of BEHAVE is that it was developed to 

produce information on the intensity and rate of spread 

of point source fires. It does not take into consideration 

changes in topography, weather, or the influences of the 

fire itself on its environment that can affect observed 

fire behavior. The original 13 models are not dynamic 

but some of the new models have been designed that 

way. Currently, BEHAVE is intended to be used by 

an experienced fire manager who can combine the 

outputs with experience to develop an expectation of fire 

behavior.

Fire Area Simulator

The Fire Area Simulator, FARSITE (Finney 1998), is 

a GIS-based fire growth model originally developed 

for planning and management of prescribed natural 

fires. Its use has since expanded to suppression efforts 

for wildfires, evaluating fuel treatments (Finney 2001; 

Stephens 1998; Stratton 200;, van Wagtendonk 1996), 

and reconstructing past fires (Duncan and Schmalzer 

2004). Developed in the Western United States, the 

model has been validated on fires in Yosemite, Sequoia, 

and Glacier National Parks (Finney 1993; Finney and 

Ryan 1995). While the use of this model is growing in 

the Southwest, Midwest, Florida, and in other countries, 

it has received little attention in the Eastern United 

States. FARSITE uses the same spread and intensity 

models as BehavePlus and, therefore, is subject to some 

of the same limitations. It requires a minimum of five 

raster layers to generate simulations: elevation, aspect, 

slope, fuel model, and canopy cover. The landscape 

data (elevation, slope, and aspect) are used for making 

adiabatic adjustments for temperature and humidity as 

well as for computing slope effects on fire spread and 

solar radiation effects on fuel moisture.

Phillips and others (2005) tested FARSITE on study 

sites in the Southern Appalachian Mountains that had 

been treated with mechanical fuel reduction (chainsaw 

felling of small trees and shrubs) or left untreated. 

They used measurements from systematically placed 

temperature sensors during prescribed burns to recreate 

test fires and compared fire behavior simulated by 

FARSITE with observed behavior. Initial simulations 

using default settings resulted in overpredictions for all 

fire behavior fuel models (FBFM) because the standard 
9	 Fire 2 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

fuel models estimate fire behavior during the western fire 

season when fuel moisture contents are low (Anderson 

1982). Actual fuel moisture values collected prior to 

burning were input into FARSITE, and subsequent 

simulations underpredicted fire spread for FBFM 9 

and FBFM 11 but still overpredicted spread for FBFM 

6. After adjustments, the rate of spread, flame length, 

and fire intensity for FBFM 9 appeared to adequately 

represent fire behavior in the leaf litter of oak-hickory 

(Carya sp.) forests. FBFM 11 seemed appropriate for 

modeling the mechanical treatment. For FBFM 6, 

adjustments allowed realistic rate of spread. However, 

the other variables of flame length and fire intensity 

were excessive, with predicted flame lengths of up to 20 

m and fire intensities up to 27,000 kW/m, which were 

not observed during the prescribed burn. The authors 

suggested that FARSITE should be viewed as an option 

for fire modeling in the Southern Appalachians, but that 

work is needed on developing fuel models that better 

represent existing conditions of fuels in this region. In 

particular, fuel moisture the and presence of ericaceous 

shrubs presented difficulties for simulations. New fuel 

models introduced by Scott and Burgan (2005) were not 

incorporated into FARSITE at the time of this test.

First Order Fire Effects Model

FOFEM, the First Order Fire Effects Model, has been 

under development by the USDA Forest Service’s Fire 

Laboratory at Missoula, Montana. It has been used 

widely by fire and land managers across the United 

States and has been endorsed by the National Wildfire 

Coordinating Group. FOFEM synthesizes the results 

of many fire effects studies into one computer program 

that can be easily and quickly used by novice or expert 

resource managers. Currently, FOFEM (v. 5.2) provides 

quantitative fire effects predictions for tree mortality, 

fuel consumption, smoke production, and soil heating 

from prescribed fires and wildfires. Although still under 

development, FOFEM provides the framework for 

predicting fire effects in most forest cover types listed by 

Eyre (1980) in the United States (http://fire.org/index.

php?option=com_content&task=view&id=57&Itemid=

31).

Tests of FOFEM for eastern hardwood systems have 

not been published, though independent use of the 

program suggests that mortality and litter consumption 

are highly overpredicted and that duff consumption 

is underpredicted. Although the scope of FOFEM is 

intended to be national, most of the models within 

FOFEM currently are based on western conifer forests. 

Mortality predictions are fitted to bark thickness and 

percent crown scorch relationships described by Ryan 

and Reinhardt (1988) for western conifers. Studies in the 

Southeast indicate that crown scorch is a poor predictor 

of mortality of southern pines (Wade and Johansen 

1986; Waldrop and Lloyd 1988) and is not relevant 

to mortality prediction for dormant-season burns in 

eastern hardwoods (Yaussy and others 2004; Waldrop 

and Mohr 2005). Litter consumption is assumed to 

be 100 percent in FOFEM; this does not reflect the 

patchy nature of burns in eastern hardwood systems 

(Waldrop and others 2004). FOFEM uses models of 

duff consumption developed by Hough (1978) for 

southern pine communities but no models are available 

for eastern hardwoods. The Joint Fire Science Program 

recently funded a proposal to modify FOFEM for use in 

the Southeastern Coastal Plain (unpublished). However, 

calibration for use in eastern hardwoods has not been 

scheduled.

The Landscape Fire and Resource 

Management Planning Tools Project

The Landscape Fire and Resource Management Planning 

Tools Project, or LANDFIRE Project, was initiated by 

federal land agencies that asked principal investigators 

to develop maps needed to prioritize areas for hazardous 

fuel reduction. The objective of the project is to 

provide the spatial data and predictive models needed 

by land and fire managers to prioritize, evaluate, plan, 

complete, and monitor fuel treatment and restoration 

projects essential to achieving the goals targeted in the 

National Fire Plan. This project will generate consistent, 

comprehensive maps and data describing vegetation, 

fire, and fuel characteristics across the United States. The 

consistency of LANDFIRE methods ensures that data 

will be nationally relevant, while a 30-m grid resolution 

assures that data can be locally applicable. LANDFIRE 

is a multiagency project among the USDA, USDI, and 

The Nature Conservancy. The principal investigators are 

located at the Rocky Mountain Research Station’s Fire 

Sciences Laboratory (MFSL) and the USGS National 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 93

Center for Earth Resources Observation and Science 

(EROS), in Sioux Falls, South Dakota) (http://www.

landfire.gov/index.html).

The first step began with the selection of two prototype 

areas with which to develop the best methods for 

mapping the rest of the country (Schmidt and others 

2002). Selected areas included the central Rockies of 

Utah and the Central Rockies of Montana, and northcentral Idaho. Protocols for producing comprehensive 

digital maps of current vegetation composition and 

condition, wildland fuel, historical fire regimes, and 

fire-regime condition class were developed using the 

Utah study area as a test case. Lessons learned during 

this process were applied to the study area in Montana 

and Idaho and completed in May 2005 (Keane and 

others 2002). The Wildland Fire Leadership Council 

chartered national implementation of LANDFIRE in 

October 2003. A national organizational structure was 

developed in addition to a project plan and associated 

schedule. The LANDFIRE technical teams at MFSL, 

EROS, and The Nature Conservancy have developed 

strategies and procedures for building on the approaches 

used in the LANDFIRE Prototype. National delivery of 

LANDFIRE products will occur incrementally over the 

next 5 years as follows: Western United States in 2006; 

Eastern United States in 2008; and Alaska/Hawaii in 

2009 (http://www.landfire.gov/ABOUT_Milestones3.

html).

LANDFIRE includes a national Rapid Assessment 

component which will deliver a “first pass,” coarse- to 

midscale assessment of fire regime conditions that is 

moderately accurate but quickly delivered. The Rapid 

Assessment will map and model potential natural 

vegetation, historical fire regimes, and fire regime 

condition class via existing datasets and expert opinion. 

The Rapid Assessment completed 12 workshops to 

create succession models and mapping rules for potential 

natural vegetation groups across the country. Participants 

included land managers and scientists from numerous 

organizations. Succession models were developed in the 

workshops for numerous biophysical settings defined as 

communities of vegetation that persist for long periods 

at landscape scales under natural disturbance regimes. 

Vegetation models currently are undergoing scientific 

and technical review (http://www.landfire.gov/ABOUT_

Milestones2.html).

Photo Series for Major Natural Fuel 

Types of the United States

Most managers have little available fuels data of the 

extent, detail, or resolution needed for fire behavior and 

effects prediction. A sequence of photos called a “photo 

series” provides a quick, easy means for quantifying 

and describing existing fuel properties for selected sites 

across a landscape. To measure fuels in the field, the user 

matches observed field conditions to the fuels shown in 

one of the photographs. Measurements of fuels given for 

each photograph can be used as a reasonable estimate 

for that area. Photo series are available for western 

systems but few exist for the East (Wade and others 

1993; Scholl and Waldrop 1999). Only two have been 

completed for the eastern hardwood region (Wilcox 

and others 1982; Sanders and Van Lear 1988). The Fire 

and Environmental Research Applications (FERA) team 

from the Pacific Northwest Research Station initiated 

a nationwide effort to produce photo series in major 

natural fuel types in 1995. The work was conducted in 

three phases. Phase I was commissioned by the USDI 

and covered 18 fuelbed types that were published in six 

volumes. Represented fuelbeds were western types with 

the exception of longleaf pine (Pinus palustris Mill.), 

pocosin, and marshgrass types in the Southeast. Phase II 

was funded by the Joint Fire Science Program in 1998. 

Eastern types included sand hill pines, hardwood/conifer 

mixtures, marshgrass, and pine/palmetto. Hardwood/

conifer sites were measured in north Georgia and 

oak/hickory sites were measured in Kentucky and Ohio. 

Phase III was funded by the Joint Fire Science Program 

and the National Fire Plan. Work began in 2001. 

Eastern types included mixed hardwoods, cutover mixed 

hardwoods, pitch pine, and additional sand hill pines. 

Publications and additional information are available on 

the FERA website: http://www.fs.fed.us/pnw/fera/.

An interesting development in newer photo series is 

the digital photo series. The Natural Fuels Photo Series 

is designed for field use and is a source of high quality 

fuels data for a variety of forest and range ecosystems 

throughout the United States. These data were developed 

for on-the-ground assessments and are not fully utilized 
94	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

in the planning environment. Technological advances 

since the inception of the original Photo Series projects 

coupled with development of new fire- and natural 

resource-based software applications highlight the need 

to bring the Photo Series concept into the electronic age. 

The Digital Photo Series will be a software application 

that will include a fuels database with a user-friendly 

interface that will leverage the already high value of the 

Photo Series data. The Digital Photo Series is nearing 

completion (http://www.fs.fed.us/pnw/fera/).

FIA National Assessment

The Forest Inventory and Analysis program (FIA) of 

the Forest Service has been monitoring forest resources 

for 75 years and represents the most comprehensive, 

consistent, and current assessment of U.S. forests 

available. FIA added measurements of fuel loading 

to a subset of its permanent inventory plots in 2001. 

The inventory, known as the Down Woody Materials 

(DWM) Indicator, has been implemented in 38 states 

to provide a regional-scale estimation of fuel complexes. 

The process measures 1-, 10-, 100-, and 1,000-hr fuels 

annually using planar intercepts, fixed radius sampling, 

and point sampling. The goal of the project is to provide 

a seamless link to fire and fuel models and to serve as a 

robust source of fuels data for national and regional fire 

scientists and managers (Woodall 2003). Fuel profiles 

are available for Minnesota, South Carolina, and the 

North Central States at http://www.ncrs.fs.fed.us/4801/

national-programs/indicators/dwm/.

Chojnacky and others (2004) examined data on DWM 

collected in 2001 by FIA on plots in several states. 

DWM data from 778 plots were used to compute 

biomass for coarse woody material, fine woody material, 

litter, duff, and shrub/herb cover. Regression equations 

were used to predict DWM components for extension 

to FIA’s more intensive plot network. Seven regression 

equations were applied to the FIA data to create maps of 

DWM biomass. General trends showed that woody fuels 

and duff were most heavily loaded in the northern states 

and decreased from north to south. The litter layer and 

the shrub and herb layer followed an opposite pattern 

with heavy abundance in the south and decreasing to 

the north. The authors suggested the patterns were due 

to differences in climate and decomposition and that 

these patterns could be used to assess regional fire-fuels 

conditions.

Regional Projects with Hardwood 

Fuel Components

Hardwood fuels have been measured in numerous 

individual studies conducted for various purposes 

throughout many eastern hardwood systems. 

Examples include low-quality hardwood stands on the 

Cumberland Plateau (Muncy 1981; Waldrop 1996), 

pine-hardwood mixtures in the Southern Appalachians 

(Phillips and Abercrombie 1987), white pine-hardwood 

mixtures in North Carolina (Clinton and others 1998), 

central hardwoods in the Missouri Ozarks (Hartman 

2004; Kolaks and others 2003; Kolaks and others 

2004; Shang and others 2004), and mixed-oak stands 

in Central Ohio (Riccardi 2002; Sutherland and 

Hutchinson 2003). Hardwood fuels were described 

for studies of stand-replacement prescribed burning in 

the Southern Appalachians by Swift and others (1993), 

Waldrop and Brose (1999), and Welch and others 

(2000). A photo series was developed for Southern 

Appalachian pine-hardwood clearcuts by Sanders and 

Van Lear (1987). The National Fire and Fire Surrogate 

Study (NFFSS) (Youngblood and others 2005) 

has three sites in eastern hardwoods: the Ohio Hill 

Country, Southern Appalachians, and the Southeastern 

Piedmont. Fuel characteristics for these sites were 

described by Iverson and others (2003), Phillips 

and others (2005), and Waldrop and others (2004), 

respectively. The Great Smoky Mountains National Park 

is conducting a parkwide vegetation mapping project 

using photogrammetric and GIS techniques to support 

an all-taxa biodiversity inventory. As a component 

of this project, fuels are being mapped using the 13 

fuel categories described by Anderson (1982). A fuel 

module was developed by He and others (2004) for the 

LANDIS forest landscape model (Mladenoff and He 

1999) and used by Shang and others (2004) to simulate 

fuel dynamics in the Missouri Ozarks.

Each of these studies provides valuable information 

about localized fuel loads and response to one or 

more variables. However, there remains a lack of 

understanding of how hardwood fuels are distributed 

across the landscape, an important and beneficial 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 95

concept for fire management planning. Studies by 

Iverson and others (2003), Kolaks and others (2004), 

and Waldrop and others (2004) suggest that loading 

of dead and down fuels is controlled by the varying 

inputs associated with different species and productivity 

levels across the landscape and varying decomposition 

rates at different sites. At any time since disturbance, 

fuel loads are a function of the amount input from 

dying vegetation minus the amount lost from decay. 

Waldrop and others (2004) also showed that fuels can 

be distributed across the landscape by gravity or by 

cultural treatments. Several studies of fuel and landscape 

interaction are described here.

LANDIS

He and others (2004) developed the LANDIS fuel 

module to simulate fuel dynamics for the Missouri 

Ozarks. The model simulates fuel inputs and 

decomposition for each of 24 land and cover types 

over a 200-year simulation. Species input data were 

from extensive inventories measured on the Mark 

Twain National Forest in the Missouri Highlands. 

Decomposition rates were those reported by Trofymow 

and others (2002) for Canadian forests. In a test of the 

model, Shang and others (2004) described fuel buildup 

and wildfire frequency that would occur under three fuel 

management treatments. A valuable product of the work 

is a GIS map of fuel accumulation across the study area 

over time. This map will allow fire managers to better 

predict where fuel amelioration is needed. LANDIS has 

been applied and tested successfully in a number of forest 

types in Missouri, Wisconsin, California, Canada, and 

China. A current effort by researchers from Texas A&M 

University and the Forest Service is the parameterization 

of LANDIS for ridgetop pine-hardwood communities 

in the Southern Appalachians. Although untested, local 

site-specific input and decomposition rates may be 

necessary to simulate the diverse topography of those 

mountains.

FORCAT

Another model designed to study dynamics of fuel 

loading was reported by Waldrop (1996). He used the 

FORCAT model (Waldrop and others 1986) to simulate 

inputs and decomposition of coarse fuels for xeric and 

mesic forest types on the Cumberland Plateau of East 

Tennessee. FORCAT is a gap model of forest succession 

that is a member of the family of models produced by 

Botkin and others (1972) and Shugart (1984). The 

simulation was set to examine fuel loads after a major 

disturbance; in this case, clearcutting was used. A 

uniform decomposition rate of 6 percent was used for 

both sites as suggested by Harmon (1982).

The estimated fuel load immediately after clearcutting 

was 49 Mg/ha on the xeric site and 69 Mg/ha on the 

mesic site (Fig. 1). On both sites, these levels were 

higher than at any other time during the 200-year 

simulation period. Fuels decompose rapidly in clearcuts 

at a period when inputs are small. In this simulation, 

decomposition exceeded inputs through year 32. 

Between years 30 and 75 there was a rapid increase 

in fuels for both simulated sites. FORCAT predicted 

decreases in stand basal area during this period as crown 

closure occurred and several large trees began to die. 

The period of rapid accumulation on the xeric site lasted 

until the stand was about 70 years old. Fuels continued 

to accumulate but at a slower rate, to a maximum at 

year 91. For the remainder of the 200-year simulation 

period, decomposition slightly exceeded inputs and fuel 

loads decreased gradually. Tree growth on the simulated 

mesic site exceeded that on the xeric site, producing a 

more rapid rate of accumulation (Fig. 1). On this site, 

fuels accumulated rapidly from years 30 through 89. 

Between years 90 and 200, fuel loading decreased much 

Figure 1.—Fuel dynamics after clearcutting xeric and mesic 

sites (predicted by FORCAT using a 6 percent decomposition 

rate for both sites).

Years

0 50 100 150 200

Fuel Loading (Mg/ha)

0

20

40

60

80

Xeric

Mesic
96	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

more rapidly than on the xeric site. Species on the mesic 

site were longer lived than those on the xeric site and 

the trees continued to grow. Mortality was higher on 

the xeric site during this period due to moisture stress. 

Therefore, inputs were less on the mesic site than the 

xeric site. Loads of coarse woody fuel projected by 

FORCAT were similar to those reported by Muller and 

Liu (1991) for old-growth stands on the Cumberland 

Plateau.

The work of Abbott and Crossley (1982) indicates 

that decomposition rates are higher on moist sites. By 

assuming a decomposition rate of 8 percent on the mesic 

site and 6 percent on the xeric site, the difference in 

simulated fuel loads between sites was greatly reduced 

(Fig. 2). Even though loading was much higher on the 

mesic site in year 1, it decomposed to a smaller amount 

than the xeric site by year 32. By year 75, fuel loading 

again was greater on the mesic site, but beyond that 

point the lines converged. During the last 50 years of 

the simulation, fuel loads on the two simulated sites 

were nearly identical. This comparison (Fig. 2) illustrates 

the observation of Abbott and Crossley (1982) that 

differences in decomposition rates between sites can 

be more important than differences in sizes or input of 

woody debris. Although the mesic site produced far more 

biomass than the xeric site, the relatively small difference 

in decomposition rates (8 vs. 6 percent) produced similar 

fuel loading throughout the 200-year simulation. This 

study is limited in scope, but suggests that fuel loads may 

be somewhat uniform across site types.

Field Measures in the Missouri Ozarks

Kolaks and others (2003) collected fuels data in the 

Southeast Missouri Ozarks to determine whether aspect 

had an effect on fuel loading in previously undisturbed 

stands. Aspect ranges included exposed slopes (135 to 

315 degrees), ridges with no aspect, and protected slopes 

(315 to 135 degrees). Fuel measurements were made at 

15 points within each of 12 experimental units. They 

found no difference in loading of 1-, 10-, and 100-

hour fuels or vertical structure of fuels across aspects, 

suggesting that the different input and decomposition 

rates across the landscape balance the loading of these 

fuels. However, the authors found significantly higher 

loading of 1,000-hr fuels on protected slopes. They 

attributed this to lower decomposition rates on moist 

sites, but we suggest (from experience gained from the 

study described below) that this is an anomaly due to 

the limited size of the study. The study was continued 

to determine whether aspect affects fuel loading after 

thinning and prescribed burning (Kolaks and others 

2004). Both thinning and burning increased fuel 

loads across all aspects, though the increase followed 

the pattern of protected > ridge > exposed. This result 

agrees with the general pattern that moist sites produce 

greater quantities of fuel. Burns consumed fine fuels 

more than those more than 3 inches in diameter. Total 

consumption did not differ among aspects but the 

data suggest that a greater proportion of fine fuels was 

consumed on exposed slopes, likely due to differences in 

fuel moisture.

Field Measures in the Southern 

Appalachian Mountains

A large-scale study was funded by the Joint Fire Science 

Program in 2001 to develop hyperspectral image maps of 

fuel loads for study sites in Tennessee, North Carolina, 

Georgia, and South Carolina (http://jfsp.nifc.gov/JFSP_

Project_Info_4.htm). The remote sensing component 

of the study will be completed by summer 2006. The 

field component is complete and has similarities to 

the studies by Waldrop (1996) and Kolaks and others 

(2003). Preliminary analyses were reported by Brudnak 

and Waldrop (2005) and Stottlemyer and others (2005). 

Additional analyses are presented here. Figure 2.—FORCAT predictions of fuel loads by site type and 

decomposition rate.

Years

0 50 100 150 200

Fuel Loading (Mg/ha)

0

10

20

30

40

50

60

70

80

Xeric - 6% Decomposition

Mesic - 8% Decomposition
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 97

Fuels data were collected at 250 randomly selected plots 

in each of four study areas (10 square miles in size). 

Study areas are on the Chattahoochee National Forest 

in northeastern Georgia, Nantahala National Forest 

in western North Carolina, Sumter National Forest 

in northwestern South Carolina, and Great Smoky 

Mountains National Park in southeastern Tennessee. 

Plot locations were generated randomly within each 

10-square-mile study area using ArcView® GIS software 

and were stratified by slope position and aspect. Fifty 

plots each were located on middle and lower slopes on 

northeast (325 to 125 degrees) or southwest (145 to 

305 degrees) aspects. An additional 50 plots were on 

ridgetops for a total of 250 plots in each of the four 

study areas. A GPS receiver was used to locate the 

plots in the field. Additional plots were included when 

necessary to provide adequate representation of all slope 

position/aspect combinations. Methods for measuring 

dead and live fuels followed standard protocols and were 

described by Brudnak and others (2005). The resulting 

dataset had measurements from 1,008 plots.

Downed woody fuels showed few differences in fuel 

loading across aspect/slope position plots (Table 1). 

Differences were observed only in the litter layer. The 

litter on the 1,008 sample plots tended to be heaviest 

along the ridges and decreased going downhill on 

both southwest and northeast slopes, suggesting that 

decomposition exceeded leaf litter inputs on the wetter 

sites. Although this difference among site types was 

significant, the relative differences were small. There 

was approximately 8 percent less litter on northeast 

lower slopes (1.68 t/acre) than on ridges (1.83 t/acre). 

There were no significant differences among slope/aspect 

combinations for loading of 1-, 10-, 100-, and 1,000-hr 

fuels or average fuel bed depth. These are preliminary 

data because analyses of impacts such as disturbance or 

cover type are not yet complete. However, these findings 

closely agree with those of Kolaks and others (2003), 

indicating that down woody fuels tend to be uniformly 

distributed across slopes and aspects.

Another component of fuels in eastern hardwood 

systems that must be considered is live fuel cover, 

particularly from ericaceous shrubs. Waldrop and Brose 

(1999) and Phillips and others (2005) reported a strong 

relationship of fire intensity to cover of mountain laurel 

(Kalmia latifolia). Van Lear and others (2002) discussed 

the importance of rhododendron (Rhododendron sp.) 

on the ecology of the Southern Appalachians. Although 

generally moist-site species, they occasionally ignite and 

can act as vertical fuels. In this study, both mountain 

laurel and rhododendron were missing from most 

measured plots but occurred in thick clumps where they 

were found. Mountain laurel was found at all aspect/

slope position combinations but was significantly more 

abundant on southwest upper slopes (Table 1). Wildfires 

that might occur could reach dangerous intensities if 

they burned uphill on dry southwest slopes and ran into 

thickets of mountain laurel. Rhododendron also was 

Table 1.—Fuel characteristics by slope position and aspect in the Southern Appalachian 

Mountains of TN, NC, GA, and SC

Slope/Aspect Litter 1 hr 10 hr 100 hr 1,000 hr Fuel Ht Kalmia Rhododendron

--------------------------t/acre-------------------------- Inches Percent Percent

Northeast

 Lower 1.68 a1 0.32 0.91 3.8 24.0 4.3 10.6 a 37.0 c

 Upper 1.82 b 0.30 0.91 3.5 18.0 4.6 13.6 a 19.7 b

Ridge 1.83 b 0.29 1.04 4.2 16.2 4.6 13.1 a 6.1 a

Southwest

 Upper 1.75 ab 0.30 0.97 3.7 17.3 4.3 21.0 b 6.8 a

 Lower 1.70 a 0.29 0.92 3.4 18.3 4.1 15.6 a 15.4 b

1

Means followed by the same letter within a column are not significantly different at the 0.05 level.
98	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

present at all slope/aspect combinations but was more 

common at lower slope and northeast-facing plots.

Fire Behavior in Eastern Hardwoods

Prediction of fire behavior in eastern hardwood systems 

is complex and includes too many variables to describe 

in this paper. Factors such as fuel moisture, temperature, 

relative humidity, windspeed, wind direction, slope, 

and aspect contribute to this complexity in ways that 

are similar to other ecosystems across North America. 

Eastern hardwoods have several differences because most 

trees are deciduous. After leaffall, abundant sunlight 

reaches the forest floor changing the microclimate there 

differently than would be the case under a protective 

canopy of conifers. Hardwood leaves generally are flat 

and can be difficult to ignite once decomposition is 

underway. However, leaves of oaks typically curl more 

than do those of maple (Acer sp.), beech (Fagus sp.), 

and birch (Betula sp.), making hardwood fuelbeds more 

complex than those of conifers. Fires that occur in dry 

weather soon after leaffall often have a high rate of 

spread but are of low severity due to a moist duff layer.

Data and other research discussed in this paper indicate 

that dead woody fuels have a fairly uniform distribution 

across different slope positions and aspects. Although 

the reason for this is unknown, it is suggested that the 

heavy fuel production that occurs on productive sites is 

balanced by rapid decomposition on those same sites. 

Regardless, managers should be more concerned with 

live fuels, such as ericaceous shrubs, jackpots (localized 

heavy loads due to a limb or tree falling), and past 

disturbances rather than the overall loading of downed 

woody fuels. Waldrop and Brose (1999) and Waldrop 

and others (2005) compared fires of several intensities 

for successful stand replacement of Table Mountain 

pine (Pinus pungens Lamb.). They found that fires that 

did not burn into jackpots or mountain laurel remained 

relatively cool throughout a 900-acre burn. Where flames 

reached a jackpot, lower limbs were scorched and some 

tree mortality occurred. If mountain laurel cover was 

over 40 percent, flames climbed into the crowns of trees 

and consumed all of the leaves. In areas where cover of 

mountain laurel was more than 85 percent, flame heights 

were twice that of overstory trees and carried from one 

crown to the next.

Additional work is necessary to better understand the 

relationships of hardwood fuels to fire behavior. Ongoing 

studies include the NFFSS sites in Ohio, North Carolina, 

and South Carolina. Numerous temperature sensors are 

scattered throughout the sites to monitor fire behavior. 

Both groups of scientists are beginning to use sensors 

located at different heights above ground to produce a 

three-dimension view of test fires.

Studies funded by the National Fire Plan and Joint 

Fire Science Program are being conducted in Ohio to 

examine landscape-scale fire behavior. The studies entail: 

1) predicting the spatial and temporal variability in key 

drivers of fire behavior, particularly fuel moisture and 

loading, and 2) monitoring fires using both airborne 

remote sensing and in-fire monitoring technologies. The 

first part of the work will provide a better understanding 

of fuel characteristics and better predictive capabilities 

as to when and how fuels will burn (that is, their 

frontal intensity, rate of spread, residence time, fuel 

consumption, and extinction). The monitoring data 

will be used to test and calibrate fire behavior models, 

including BEHAVE and a coupled fire-atmosphere model 

(http://www.fs.fed.us/ne/delaware/4153/4153.html).

SUMMARY

Although research on fire in eastern hardwoods has 

lagged behind that of other U.S. regions, there are 

numerous ongoing studies and several prediction 

products already available. The BEHAVE system tested 

well in two hardwood systems for predicting rate of 

spread, especially when customized fuel models were 

available. FARSITE has not been used extensively in the 

region but shows promise as local fuel measurements 

become available. FOFEM includes the computer code 

to allow prediction of fire effects in eastern hardwoods. 

However, it has not been calibrated for the region. 

LANDFIRE is an aggressive nationwide effort to provide 

fuel and vegetation maps of the entire country. Products 

for eastern systems should be available in 3 years. Phase 

III of the Photo Series for Major Natural Fuel Types 

of the United States is nearing completion and should 

include fuel models for eastern hardwoods. And the 

Forest Service FIA is measuring fuels across the United 

States and providing valuable information about regional 

fuel patterns.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 99

In addition to nationwide programs, numerous local 

studies have been completed that describe local fuel 

types. Three independent studies compared hardwood 

fuel loads for different slope positions and aspects. 

All found few differences and suggested that higher 

productivity on moist sites was balanced by higher 

decomposition on those same sites. Major sources of 

fuel, and thus areas of concern for fire managers, were 

localized jackpots and presence of mountain laurel and 

rhododendron. Additional work is needed to better 

predict fuel loads throughout the eastern hardwood 

region and to model fire behavior within complex 

interactions of weather, fuel, and topography.

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1	 Fire 04 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

SOIL, FIRE, WATER, AND WIND: 

HOW THE ELEMENTS CONSPIRE IN THE FOREST CONTEXT

Ralph E.J. Boerner1

Abstract.—Reviews our current understanding of the impact of fires typical of eastern oak forests on 

soil properties, soil organisms, and water quality. Most oak ecosystem fires are dormant-season fires 

whose intensity falls at the low end of the range of wildland fires. Direct heating of the mineral soil 

generally is minor except where accumulations of woody debris smolder for lengthy periods. Considerable 

proportions of nitrogen, phosphorus, and cations may be lost during fire through a combination of 

volatilization and ash convection. Post-fire precipitation events govern the return of nutrients in ash to 

the mineral soil, and the interaction of the soil exchange capacity, geomorphology, and weather control 

the proportion of nutrients from ash that will be retained for later plant use. Exposure of mineral soil by 

fire may lead to increased sheet erosion, but soil hydrophobicity does not seem to be important in oak 

ecosystems. Nitrogen availability and organic carbon content of soils may increase after fire, though both 

appear to be lesser in magnitude and duration than in other ecosystems. Impacts on fungi, bacteria, and 

microarthropods in the mineral soil are small, whereas those in the forest floor are proportional to the 

degree of consumption and the extent of heating due to smoldering woody fuels. Microbial activity and 

microarthropod populations recover quickly except after repeated annual burning. Little is known of 

effects of fire on other groups of soil organisms (e.g., nematodes and earthworms). The geomorphology 

of much of the eastern oak region is complex and heterogeneous. In such terrain, the difficulties inherent 

in scaling up plot-based studies to land areas of management scale are considered in the context of 

demonstrated landscape-scale variations in belowground effects of fire. GIS-derived landscape-scale metrics 

can be used to help generate management-scale recommendations from smaller scale research studies.

to the use of fire by Native Americans in oak forests and 

savannas, from Massachusetts to Missouri, and from 

the early 1600’s to the early 1800’s. Some of these early 

observers even commented on the positive (Lorain 1825) 

or negative (Coxe 1794; Lorain 1825) effects of fire on 

forest soil productivity.

The key role that fire plays in the ecology of oakdominated forests was recognized by ecologists as early as 

the 1920’s and 1930’s (e.g., Daubenmire 1936; Cottam 

1949). Ironically, this is the same time period in which 

organized fire suppression policies came into force and 

became effective, at least in Ohio (Sutherland and others 

2003).

Quantitative studies of the effects of fire on the soils of 

oak forests, and on the waters draining such forests, did 

not become common until after World War II. Thus, 

current and recent studies of the effects of fire on forest 

soil and water have taken place not in the context of the 

long, continuous history of fire that preceded the 20th

century but in the artificial context of forests already 

affected by many decades of fire suppression.

INTRODUCTION

Analysis of pollen and charcoal deposits taken from 

water bodies along the plateaus of the Appalachian 

Mountains demonstrates that fire has been a frequent 

and consistent part of the forests, savannas, and 

grasslands of Eastern North America for at least the last 

4,000 years (Delcourt and Delcourt 1997). Further, 

evidence from macrofossils and pollen indicates that 

forests dominated by fire-dependent trees such as oak 

(Quercus) and pine (Pinus) have covered much of Eastern 

North America for at least 10,000 years (Delcourt and 

Delcourt 1987). Thus, the history of fire in eastern 

forests precedes the development of significant human 

populations in the region.

During and after European settlement, many observers 

noted the use of fire by Native Americans. Whitney 

(1994) summarized more than 20 historical references 

1

Department of Evolution, Ecology, and Organismal Biology, 

The Ohio State University, Aronoff Laboratory, 318 West 

12th Avenue, Columbus, OH 43210, 614-292-6179, email: 

boerner.1@osu.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 105

FIRES IN OAK FORESTS

The term “forest fire” conjures images of fires in 

western pine forests, with flames rising 30 meters or 

more, trees exploding, and fire storms carrying burning 

brands for miles. Although such intense fires can occur 

in our region, especially during lengthy droughts 

in forest types with a significant pine component 

(Forman and Boerner 1981), such intense, standreplacing fires are uncommon in oak forests. Most 

fires in the oak forests of the central hardwoods occur 

during the dormant seasons (spring and autumn). 

Such fires typically move along the forest understory 

without climbing into the canopy (Sutherland and 

others 2003).

Fire has both immediate, direct effects as well as 

delayed effects on soil, soil organisms, and water that 

emerge in the days, weeks, and months after fire (Fig. 

1). The degree to which those effects are felt is, in turn, 

regulated by weather, by the structure of the landscape, 

and by the legacies of past management, fire history, and 

land use (Fig. 1). In reviewing the available information 

on these topics I will rely where possible on studies from 

eastern oak forest ecosystems, and will supplement with 

studies from other temperate ecosystems (particularly 

pine forests and shrublands) only where there is 

insufficient direct evidence from eastern oak forests. 

Those interested in reviews of fire effects on ecosystems 

in general may consult recent, broadly based reviews, 

such as those of Neary and others (1999), Ice and others 

(2004) and Certini (2005).

SOIL HEATING

As a generalization, surface fires moving across the floor 

of a forest do not present a strong potential for severe 

soil heating because only a small percentage of the heat 

generated by the fire is partitioned downward into the 

soil. Further, mineral soil is a poor conductor of heat, 

especially when the soil is relatively dry (Raison 1979).

How much the temperature of the mineral soil increases 

during fires is the key to understanding the impact of 

the fire on soil organisms. Temperatures of 70°C for 

as little at 10 minutes will kill almost all fungi and 

soil microfauna, and also a significant proportion of 

soil bacteria (Lawrence 1956). Temperatures of 60°C 

are considered lethal to plant roots (Steward and 

others 1990) and temperatures above 60 °C denature 

enzyme systems of metabolically active cells of most 

microorganisms.

In areas where a distinct humus layer is absent or 

where that forest floor layer does not begin to smolder, 

insufficient heat is transferred downward into the 

mineral soil for temperatures to rise above ~60°C at 

depths below several centimeters (Anson and Gill 

1976; Steward and others 1990). Saa and others 

(1993) measured soil temperatures at a depth of 5 cm 

in the mineral soil during surface fires in pine forests 

and shrublands in Spain. They did not record soil 

temperatures ≥ 50°C either during or after fire. Similarly, 

Hayward (1938) recorded soil temperatures during a fire 

in a longleaf pine (Pinus palustris Mill.) forest in North 

Carolina and found that temperatures at 2.5 cm into 

the mineral soil did not exceed 40°C at any time during 

or after the fire. Thus, fires moving across the forest 

floor are unlikely to produce increases in mineral soil 

temperatures sufficient to cause significant soil organism 

mortality.

One situation in which temperatures may rise 

sufficiently to induce considerable mortality is when 

localized concentrations of organic materials, either 

Figure 1.—Schematic of immediate fire effects and post-fire 

emergent effects on soil physical, chemical, and biological 

properties, with regulating factors indicated at left.
1	 Fire 06 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

as woody debris (Miller and others 1955; Busse and 

others 2004) or as a particularly thick, dry humus layer 

(Miyanishi 2001), smolder in place for an extended 

period. Miller and others (1955) monitored surface 

and mineral soil temperatures during and after a fire in 

a New Zealand shrubland. aThere was no significant 

change in temperature at a depth of 5 or 10 cm in the 

mineral soil under this fire as it moved across the site. 

However, in areas where localized wood heaps burned for 

several hours, sensors recorded maximum temperatures 

of 100°C at 5 cm and 60°C at 10 cm into the mineral 

soil. Similarly, temperatures of 100 to 200°C have been 

recorded under smoldering Arctostaphylos residues at 

a depth of of 7.5 to 12.5 cm in a mixed-conifer forest 

in California (Busse and others 2004). Thus, localized 

heavy fuel accumulations such as slash piles from 

harvesting or detrital accumulations from storm damage 

present a situation in which direct heating effects may 

have significant negative effects on soil biota, including 

plant roots.

VOLATILIZATION AND ASH 

CONVECTION

Two pathways by which nutrients may be lost from 

an oak forest during and shortly after fire are direct 

volatilization (conversion of elements in fuel to gas 

phase) and ash convection (conversion of elements to 

solid, inorganic ash subject to wind action). Which 

nutrients are subject to being volatilized depends on 

the element’s chemical properties, the properties of the 

compound in which that element is found, and the 

behavior of the fire. Similarly, losses via ash convection 

depend on the local weather conditions, the behavior 

of the fire, and the characteristics of the fuels being 

combusted.

Under controlled laboratory conditions, organic 

compounds containing nitrogen (N), phosphorus 

(P), and sulfur (S) often begin to volatilize when 

temperatures approach or exceed 200°C (Table 1). The 

temperatures reached in the fuelbed (~650°C) during 

flaming or smoldering after the flames have passed are 

considerably higher than that critical temperature. This 

helps explain why the amount N lost by volatilization is 

so closely dependent on the amount of fuel consumed 

(e.g., Raison and others 1985).

One of the few estimates of the loss of N via this 

pathway in oak forests is the report of Hubbard and 

others (2003) of losses of 55 kg N/ha from litter + wood 

and 6 kg N/ha from humus as the result of prescribed 

fires in oak-pine stands in Georgia and Tennessee. They 

indicated that losses of this magnitude could be replaced 

by atmospheric deposition of N in a relatively short time. 

One also can gain perspective on the potential 

importance of this pathway of N loss from estimates 

generated in other temperate ecosystems dominated by 

woody plants. Losses of N from vegetation and detritus 

to the atmosphere have been estimated at 70 percent in 

a North Carolina longleaf pine savanna (Christensen 

1977), 33 percent in a mixed pine plantation in South 

Carolina (Wells 1971), 39 percent in a mixed conifer 

forest in Washington (Grier 1975), and 39 percent in a 

chaparral shrubland in California (DeBano and Conrad 

1978). In general, losses of N to the atmosphere are 

proportional to fuel mass loss during fire (Raison and 

others 1985). 

Compared to N, base cations such as calcium (Ca), 

magnesium (Mg), and potassium (K) and inorganic 

forms of P have considerably higher laboratory 

volatilization temperatures (Table 1). Nonetheless, 

losses of cations and P from vegetation and detritus 

during a fire can be significant. Estimates of Ca loss 

range from 11 percent in a Washington mixed conifer 

forest (Grier 1975) to 15 percent in a longleaf pine 

savanna (Christensen 1977), to 17 percent in a mixedeucalypt forest (Harwood and Jackson 1975). The same 

three studies report losses of K ranging from 9 to 46 

percent and losses of Mg ranging from 13 to 17 percent. 

Similarly, estimates of the loss of P through this pathway 

range from 10 percent in a mixed-eucalypt forest in 

Tasmania (Harwood and Jackson 1975) to 46 percent in 

the longleaf pine site described by Christensen (1977).

The degree of combustion of the fuels and the resultant 

nature of the particulate materials available for transport 

can influence the relative importance of volatilization 

and ash convection as pathways for nutrient export. 

Raison and others (1985) determined that low intensity 

fires in three subalpine Eucalyptus forests resulted a ratio 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 107

of 57 percent particulate to 43 percent nonparticulate 

loss of P when combustion produced black ash, 

and a ratio of 73 percent particulate to 27 percent 

nonparticulate loss when combustion proceeded all the 

way to very fine grey ash.

Although small losses of cations via direct volatilization 

may occur even in low intensity fires (e.g., Raison and 

others 1985), losses from the ecosystem of P and cations 

of the magnitudes listed also must represent exports 

of ash and other particulate combustion products 

due to the action of either ambient wind or air drawn 

(entrained) into the combustion zone because of the 

buoyancy of hot flame gases. Fire intensity (the rate of 

heat release from flaming per unit length of fire line, 

kW/m) determines both flame length and the velocities 

of air entrainment. Thus, it would be expected that 

losses of nutrients by ash convection would increase with 

fire intensity, a pattern suggested by Raison and others 

(1985). In addition, fire intensity tends to increase when 

fires burn in wind, so wind and fire intensity often may 

act together to increase nutrient losses in ash. Given the 

strong spatial heterogeneity in fuel consumption and 

intensity typical of most oak ecosystem fires, there likely 

are large spatial differences in nutrient losses either from 

volatilization or ash convection. Thus, one can generalize 

that losses of nutrients to the atmosphere may result 

in large losses of nutrients from the ecosystem, what 

proportion of the ecosystem nutrient capital is lost and 

via which atmospheric pathway depends on the element 

of interest, its biochemical context, the local weather 

conditions, and the behavior of the fire.

ASH DEPOSITION AND SOIL 

NUTRIENT AVAILABILITY

The material that remains after fire has consumed 

part or all of the fuel it can access is a combination of 

patches of uncombusted material, partially combusted 

woody material, and ash, the end product of complete 

combustion of the carbon (C) skeleton of the fuel 

materials. The inorganic materials that comprise the 

ash/partially combusted materials and whatever soluble 

organic compounds are present in the latter are all easily 

dissolved by rainfall. Whether these materials remain 

in the mineral soil or are lost from the system through 

leaching or overland runoff depends on several factors, 

most notably weather patterns, topography, and the 

properties of the mineral soil at that site.

During the first month after an early spring fire in a 

South Carolina pine plantation complex, approximately 

80 mm of rainfall resulted in ~70 percent of the Ca and 

K in the ash being dissolved and transported downward 

into the soil (Lewis 1974). Similarly, in a study of 

post-fire nutrient dynamics in a mixed conifer forest 

in Washington, Grier (1975) observed that 90 percent 

or more of the ~150 kg Ca/ha, 50 kg K/ha, and 85 kg 

Mg/ha present in ash were dissolved and transported 

during the first precipitation event after the fire: 67 

cm of snowmelt. In both of these cases, there was little 

actual loss from the ecosystem as the vast majority of the 

nutrients dissolved from the ash remained in the mineral 

soil.

Table 1.—Estimates of temperatures at which various 

elements/compounds of ecological importance 

volatilize under laboratory conditions; elements/

compounds are listed in order of increasing 

minimum volatilization temperature

Element Form Volatilization

°C

N organic forms 175-200a

N fumaric acid dinitrile 189b

N alanine <200b

N adenine <200b

P phosphoric acid 203b

P phosphobenzoic acid 249b

N aminobenzoic acid 249b

P organic forms 340-360a

S organic forms 375a

K inorganic forms 550a

P inorganic forms 770a

K base metal 774b

Ca CaCO

3

825-898b

Mg MgCO3900b

Mg base metal 1107b

P potassium 

hexametaphosphate

1320b

Ca base metal 1487b

a

Agee (1993). b

Weast (1969).
1	 Fire 08 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

The impact of ash deposition and dissolution on soil 

nutrient status is most obvious when fire has been 

both frequent and concentrated in space. For example, 

in their study of an oak forest site in Pennsylvania 

that had been used as a charcoal hearth from 1771 to 

1884, Mikan and Abrams (1995) reported that Ca and 

Mg availability in the soils under the former hearth 

were 12.1 and 4.8-fold greater, respectively, than in 

the surrounding forest. Thus, repeated deposition of 

concentrated ash caused changes in nutrient status that 

were still apparent more than a century after the last 

hearth fire. 

A similar example is from the study of the effects of 

burning concentrated slash piles (i.e., jackpot fires) after 

logging in an English oak forest (Jalaluddin 1969). In 

this study, the slash was piled in areas of < 2.0 m in 

diameter and burned for nearly 3 hours. Samples taken 

several days after the fires indicated that dissolution 

of the resulting ash had increased soil pH from 6.0 to 

9.0. However, within 6 months, continued leaching of 

the soil resulted in the pH returning to ~ 6.0. Thus, 

the effects of fire on nutrient status are sensitive to the 

amount of ash deposited in a fire, the number of fires on 

that site, and the intensity/duration of the leaching of 

the site by precipitation following ash deposition.

After a single fire in a pine-oak ecosystem in Kentucky, 

Blankenship and Arthur (1999) reported that soil pH 

had increased by 0.2 to 0.3 unit, demonstrating the 

effect of a single fire and subsequent ash dissolution on 

soil base status. After two or four fires in a mixed-oak 

ecosystems in Ohio, Boerner and others (2004) reported 

fire-induced, significant and persistent increases in soil 

pH of 0.2 to 0.6 unit. This study also reported increases 

in extractable Ca2+, reductions in extractable aluminum 

(Al3+), and in increased molar Ca:Al ratios, though 

these differences were statistically significant only in the 

more nutrient-poor sites (Boerner and others 2004). In 

sites with significant limestone influence in the soils, 

changes in base cation status were transitory. Similarly, 

Knoepp and others (2004) observed increased soil 

pH, exchangeable Ca, and exchangeable Mg after site 

preparation (fell-and-burn) burning in western North 

Carolina.

Although most studies of the effects of fire on nutrient 

status focus on a single fire or several fires over a decade 

or less, two longer term experiments in oak forests 

may help shed light on chronic effects of burning on 

oak forest soils. Eivasi and Bayan (1996) assessed the 

nutrient status of soils in oak-hickory flatwoods in 

Missouri that had been subjected to annual or periodic 

(~ 4 year interval) burning for 40 years. They found 

no significant effects of fire at either frequency on soil 

pH, Ca, K, or Al. They did report that available P was 

reduced to 24 percent of controls by annual burning and 

to 35 percent of controls by periodic burning. Thor and 

Nichols (1973) reported a similar lack of change in pH 

from eight annual or two periodic burns in oak stands 

on the Highland Rim region of Tennessee. DeSelm and 

others (1991) resampled the Thor and Nichols (1973) 

sites after 27 years of annual or periodic burning, and 

again reported that burning had no significant effect on 

soil pH or the availability of K or P. Thus, although the 

effect of lengthy periods of burning had no effect on 

base-cation availability in either ecosystem, the longterm effect of low-intensity fire on P availability remains 

unclear.

No aspect of the belowground responses to fire has been 

more intensively studied than the availability of N in the 

soil. This is partially the result of the generally accepted 

view that most ecosystems are N-limited. Whether 

eastern oak ecosystems are still N-limited in 2005 given 

the chronic deposition of N from fossil fuel combustion 

and agricultural sources is an open question and beyond 

the scope of this review.

Wan and others (2001) performed a meta-analysis of 

the effects of fire on soil N availability and concluded 

that, over all ecosystem types, inorganic N availability is 

increased by fire. Ammonium (NH4

+

) availability peaks 

soon after fire but returns to pre-fire levels in less than 

a year. The pulse of NH4

+

 is due to a combination of 

liberation from organic matter degraded during the fire, 

activity of heterotrophic soil biota, and N-fixation by 

symbionts of newly colonizing plants. Nitrate (NO3

-

) 

availability peaks some months later (generally 6 to 12

months after fire), and is the result of enhanced NH4

+

availability and increased activity of nitrifying bacteria. 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 109

However, when the results were stratified by fire type or 

intensity, Wan and others (2001) concluded that high 

intensity wildfires and slash fires resulted in increased 

NO

3

-

 and NH

4

+ whereas prescribed burning did not. 

As the majority of fires in eastern oak forests may be 

more like prescribed fires than wildfires or slash fires in 

grasslands, shrublands, or coniferous forests, one might 

speculate that the conclusions of Wan and others (2001) 

would have been different had their analysis been limited 

to ecosystems with a significant oak component.

Soil organic matter is a key part of a forested ecosystem. 

This material is an essential store of nutrients (especially 

N and C), is essential to exchange processes that regulate 

the availability of Ca, Mg, S, P, and K to plants and 

microbes, helps stabilize soil structure by cementing soil 

particles into stable aggregates, and insulates the soil 

against changes in microclimate. Enhancement of soil 

organic matter development in forests also is a pathway 

that is being examined as a possible mechanism for 

ameliorating the effects of CO2 released by fossil fuel 

combustion.

Johnson and Curtis (2001) performed an extensive 

meta-analysis of the effects of disturbance on soil organic 

matter in forests. They concluded that ecosystems that 

had experienced fire approximately 10 years earlier had 

an average gain in soil organic C of approximately 8 

percent. They attributed this change to a combination 

of factors, including infiltration into the mineral soil 

of organic matter from partially combusted fuels, 

conversion of labile (easily decomposed) organic matter 

into recalcitrant (stable) organic matter, and the effects of 

the colonization of burned areas by plants which harbor 

N-fixing symbionts.

Despite a rich literature from coniferous ecosystems 

that demonstrates significant loss of soil organic matter 

during and after fire, few studies have demonstrated 

major changes in soil organic-matter content following 

fire in eastern oak ecosystems. Knighton (1977) observed 

no significant change in soil organic C following one to 

three fires in Wisconsin oak forests, and Knoepp and 

others (2004) reported no significant change in soil 

C over 5 years following site preparation burning in 

oak-pine stands in North Carolina. We observed only 

slight changes in soil organic matter in Ohio oak-hickory 

sites subjected to one to four prescribed fires (Boerner 

and others 2000b; Boerner and Brinkman 2004). It 

is important to note that the lack of change in soil 

organic matter after fire may be specific to low-intensity, 

dormant-season fire regimes. For example, soil organic 

matter was reduced by as much as 50 percent following 

intense wildfire in an Israeli pine-oak (Pinus halepensis 

Mill-Quercus calliprinos Webb) ecosystem (Kutiel and 

Naveh 1987).

Results from single fires or from studies of several fires 

to the longer term are projected with considerable 

uncertainly. And, this is exacerbated by somewhat 

conflicting results from the two longer term studies of 

fire in oak forests that do exist. Eivasi and Bayan (1996) 

could not detect significant changes in soil organic-matter 

content in the soils of a Missouri oak flatwoods ecosystem 

even after more than 40 years of annual or periodic 

burning. By contrast, Philips and others (2000) reported 

significant reductions in A horizon organic-matter 

concentration and content after 35 years of annual fires 

(but not periodic fires) in a Tennessee oak forest.

Some of the material deposited on the ground surface 

after a fire is partially combusted, charred woody 

material. The blackened materials that comprise a 

considerable portion of this charred fuel have been 

termed “black carbon,” and have become a focus of 

research on organic matter changes after fire (Certini 

2005). Black carbon can comprise as much as 40 

percent of the soil organic matter in ecosystems exposed 

to frequent fire (Ponomarenko and Anderson 2001), 

and this material may have sorptive properties that 

are important in regulating soil solution chemistry 

and biochemistry for some time after fire (Wardle and 

others 1998). However, no estimates of the abundance 

or importance of black carbon in oak forests have been 

published.

EXPOSURE, EROSION, AND 

HYDROLOGY

Changes in the soil water and hydrological regimes of an 

ecosystem typically are proportional to the proportion 

of the litter, humus, coarse woody debris, and woody 

stems that are consumed during the fire. Fires that 
11	 Fire 0 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

consume most or all of the forest floor and root systems 

may produce major changes in watershed hydrology. 

For example, Folliot and Neary (2003) reported an 

increase of 90 percent in watershed output after a highintensity fire in a ponderosa pine (Pinus ponderosa Laws.) 

ecosystem in Arizona. By contrast, low-intensity fires 

that do not consume the entire forest floor and/or fires 

that affect only part of a watershed may not produce 

hydrological changes that can be resolved by typical 

monitoring methods (Bethlalmy 1974).

The volume of water leaving a forested watershed in 

streamflow and ground water recharge is a function of a 

range of factors, including interception of precipitation 

by the canopy and subsequent evaporation, transpiration 

by plants, evaporation from the ground surface, overland 

runoff, water storage in the soil, and transport of water 

through the soil profile to groundwater or surface 

waters. As low-intensity, dormant-season fires in eastern 

oak forests typically do not have a major impact on 

canopy structure and understory vegetation grows/

regrows rapidly after fire, neither canopy interception/

evaporation nor transpiration likely will be affected 

strongly by fire. Evaporation from the blackened soil 

surface may be enhanced after fire but such an effect 

likely will be limited to the period before the canopy 

leafout and understory development in the spring 

(Iverson and Hutchinson 2002).

Fires that consume the entire forest floor also may 

affect the water-holding capacity of the top several 

centimeters of the soil through changes in soil texture 

and structure (Austin and Baisinger 1955). Severe 

heating may cause clay-size particles to fuse into larger 

particles, thereby affecting soil texture and porosity; 

however, the intensity necessary to produce such effects 

is not typical of oak ecosystem fires. Changes in surface 

soil permeability or porosity also may occur as the result 

of the volatilization of organic compounds that had 

served to cement soil aggregates (Dyrness and Youngberg 

1957). This can result in reduced porosity as impacted 

aggregates crumble and plug large soil pores (Moehring 

and others 1966). Severe and/or repeated burning also 

has the potential to increase soil bulk density, though 

such effects typically are less evident after low intensity, 

periodic fires (Moehring and others 1966; Agee 1973).

Whether water impacting on the forest floor/soil surface 

percolates into the soil or runs off over the surface 

depends not just on the permeability of the soil but also 

on whether water is free to move downward through 

the soil. In ecosystems where coarse-textured soils are 

overlain by resinous forest floor materials, combustion of 

the forest floor can cause volatilization of hydrophobic 

organic compounds. If there is significant downward 

transport of those volatilized materials, they may 

condense as hydrophobic coatings on soil aggregates, 

resulting in the development of what has been termed 

a hydrophobic or water-repellant layer (Agee 1973). 

Significant hydrophobicity has been observed after fire in 

coniferous forests and chaparral (e.g., DeBano and others 

1970), and once developed can result in significantly 

increased erosion, as water that would normally 

percolate vertically through the profile is instead routed 

downslope through the surface soils along the waterrepellant layer (Agee 1973). Although examples of 

fire producing nonwettable soil layers are common in 

coniferous ecosystems to the west and north of the oak 

forest region, I have found no reports of hydrophobicity 

developing after fires in oak forests.

As regulation of water flow and water quality are among 

the most important ecosystem services we expect forested 

watersheds to supply, the issue of the effect of fire on 

water quality merits consideration. Again, in ecosystems 

subject to large, intense fires, water quality may be 

affected greatly by surface runoff and particulate erosion. 

However, in ecosystems such as the eastern oak forests, 

fires do not typically produce effects on the watershed 

that are likely to produce such large or chronic changes 

in erosion and runoff. Instead, leaching of soluble 

materials from the ash and partially combusted fuels 

through the soil profile and into ground water or water 

bodies is of greater concern.

Knighton (1977) analyzed the effects of zero to three 

annual burns on the quality of water leaving an oakhickory forest in central Wisconsin. He reported 

solution concentrations of Ca2+ and Mg2+ to be 

significantly greater in leachate below the burned areas 

than in leachate from unburned controls. The solution 

concentrations of NO

3

-

 and PO

4

3- also were greater in 

leachate from burned sites, and the degree of enrichment 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 111

was proportional to the number of fires a site had 

experienced. However, Knighton (1977) pointed out 

that the increases in N and P concentration with fire 

frequency were not statistically significant due to highamong sample variability. Similarly, Lewis (1974) noted 

increases of 50 to 125 percent in concentrations of Ca2+

and K+ (but little change in NO3

-

 and PO

4

3-) in soil 

leachate after fires in South Carolina pine plantations, 

but great variation among samples resulted in the 

differences not being statistically significant. Whether 

water quality is affected by low-intensity, dormant-season 

fires is likely to be determined by the relationship among 

ash deposition, post-fire precipitation, and soil nutrient 

storage capacity. Thus, predictions of fire effects on water 

quality will be site and situation specific.

SOIL BIOTA AND ECOSYSTEM 

PROCESSES

The assemblage of animals in the forest floor and soil of 

a forested ecosystem is both highly diverse and critically 

important to ecosystem processes. These animals form 

a complex food web that is responsible for much of the 

conversion of coarse detritus produced both belowand aboveground into fine organic materials that are 

suitable for subsequent microbial decay. Without these 

micro- and mesofauna present, leaf litter decomposition 

essentially ceases (Edwards and Heath 1963).

Of the many groups of arthropods present in and on 

the forest floor and soil of forested ecosystems, the 

Collembola (springtails) and Acari (mites) are the most 

important to the decay process (see also Rieske-Kinney, 

this volume). In southern Ohio mixed-oak forests, 

three annual burns reduced the numbers of springtails 

and mites by more than 50 percent, while a single 

fire resulted in no significant change in abundance 

in either group (Dress and Boerner 2004). There are 

even suggestions in that study that conditions after 

a single fire may have supported greater densities of 

microarthropods than did those of the unburned 

controls. Metz and Farrier (1971) reported that mites 

and springtails in the forest floor decreased in abundance 

in direct relation to fire frequency in a South Carolina 

pine plantation. When fires were applied every 4 

years, there was sufficient time for recolonization and 

population growth to return numbers to pre-fire levels, 

whereas when fire was applied annually, abundances 

remained depressed over time. By contrast, Metz and 

Farrier (1971) reported no significant effect of annual or 

periodic fire on mites and springtails in the mineral soil.

As sparse as the database for microarthropods might be, 

that for other faunal groups is even more depauperate. 

Matlack (2001) reported no longterm effect of fire 

on nematode numbers or diversity in southern pine 

plantations. Bhadauria and others (2000) observed that 

annelid earthworm abundance was depressed by fire in a 

mixed oak-pine forest in northern India. However, they 

also reported that earthworm abundance returned to prefire levels over several months after fire as the result of 

migration from neighboring, unburned areas. Bhadauria 

and others (2000) suggested that post-fire conditions 

in the burned areas might be better suited for annelid 

growth than conditions present in unburned areas.

The effects of fire on the soil microbial community can 

be evaluated by examining individual species, species 

groups, or microbially mediated processes such as the 

conversion of organic, N-containing compounds to 

inorganic N as NH4

+

 (N mineralization). In a South 

Carolina pine plantation, Jorgensen and Hodges (1971) 

found that both fungal and bacterial abundance in the 

forest floor were reduced by annual fire, but abundances 

of both groups in the forest floor of plots burned on 

a 4-year cycle were greater than those of unburned 

controls (Fig. 2). The same pattern was apparent for 

bacteria in the upper 5 cm of the mineral soil, whereas 

Figure 2.—Effects of fire on soil and forest floor 

microorganisms, redrawn from Jergensen and Hodges (1971).
112 	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

fungal abundance in the mineral soil was not affected 

by fire treatment (Jorgensen and Hodges 1971). The 

notion expressed in this study that periodic fire produces 

conditions for soil microbes superior to those present 

in unburned sites parallels those from the studies of soil 

invertebrates described earlier (Bhadauria and others 

2000; Dress and Boerner 2004.

Blankenship and Arthur (1999) demonstrated a 

significant, positive effect of a single winter fire in an 

eastern Kentucky oak-pine ecosystem on bacterial 

biomass, and this was accompanied by an increase in the 

bacteria:fungi biomass ratio in this ecosystem. A similar 

shift in the bacteria:fungi ratio after a low-intensity 

slash fire was reported by Perry and others (1984) in a 

northwestern coniferous forest. As the fungi in those 

coniferous systems are responsible for producing the 

compounds that make Fe available for plant uptake 

(hydroxymate siderophores), Perry and others (1984) 

believed that there was a potential for iron deficiency 

of tree seedlings after fire, especially where pH has 

been increased. Whether such short-term effects on the 

microbial community occur broadly in oak ecosystems 

and for how long such effects might persist is unknown. 

Fonturbel and others (1995) found no chronic or lasting 

effect of prescribed fire on heterotrophic bacteria or 

filamentous fungi in a pine ecosystem in Spain, but the 

applicability of this finding to eastern oaks is uncertain. 

Eivasi and Bayan (1996) found that 40 years of annual 

or periodic burning reduced microbial biomass in a 

Missouri oak forest by 32 and 21 percent, respectively.

Mycorrhizal fungi are key symbionts for virtually all 

forest plants. There are two major groups of mycorrhizal 

fungi found in the eastern oak regions (as well as several 

more limited groups of mycorrhizae sensu latu that are 

not considered here). Ectomycorrhizal fungi (ECM) 

are a group of higher fungi (basiodiomycetes and 

ascomycetes) that form symbioses with many conifers 

(e.g. Pinus, Picea, Abies), oaks (Quercus spp.), hickories 

(Carya spp.), and beech (Fagus grandifolia Ehrh.). 

Arbuscular mycorrhizal fungi (AMF), also referred to 

as endomycorrhizae, vesicular-arbuscular mycorrhizae 

and (VAM), are a much smaller and less diverse group 

of lower fungi that form symbioses with herbaceous 

plants and with woody plants that do not form ECM 

symbioses. Both AMF and ECM are critical to plants for 

the acquisition of P, as P is relatively immobile in most 

forest soils. In addition, ECM can be important in N 

uptake and pathogen resistance.

The colonization of roots by arbuscular mycorrhizal 

fungi is sensitive to soil heating, at least under laboratory 

conditions. Colonization of plant roots by AMF was 

reduced by ~50 percent when soil temperatures exceeded 

50 to 60°C, and was reduced by 95 percent when soil 

temperature reached 90°C (Klopatek and others 1988). 

Whether this is important under field conditions is 

uncertain, as Knorr and others (2003) reported no 

significant effect of one to four fires on the abundance 

of the AMF-specific glycoprotein glomalin in oak forest 

soils in Ohio. 

Severe fires can alter ECM community structure in 

bishop pine (Pinus muricata D.Don.) stands on the 

Pacific coast (Baar and others 1999). The decrease in 

ECM abundance and diversity that takes place after fire 

in western conifer forests may occur because the bulk of 

the fungal mycelium in those ecosystems is located in 

the humus layer of the forest floor, and that layer often 

is completely lost in severe wildfires (Harvey and others 

1986). However, the applicability of this work to eastern 

oak ecosystems is uncertain. First, not all eastern oak 

forests exhibit a strongly developed humus horizon in 

the forest floor. Second, two studies in oak ecosystems 

suggest that fungi present in the mineral soil can rapidly 

recolonize the forest floor in the months after fire.

Tresner and others (1950) found that the species 

composition of fungi in a Wisconsin oak forest did not 

change with depth from the forest floor down through 

the upper mineral soil. They concluded that colonization 

of disturbed surface layers from deeper soil is possible 

without changing fungal community structure. 

Jalaluddin (1969) examined fungal dynamics in soils 

of small plots on which concentrated piles of slash had 

been burned. Fungal abundance in the center of those 

intensely burned plots was only 3, 6, and 11 percent 

of control levels at 1 week, 3 months, and 6 months 

after fire. However, at the edges of the burned plots, 

where recolonization from neighboring, unburned soils 

could occur, abundances were 17, 31, and 43 percent of 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 113

control levels at those same points in time. Thus, even 

when intense fire effectively sterilizes the mineral soil, 

thus precluding recolonization from below (sensu Tresner 

and others 1950), mycelial ingrowth from surrounding 

areas can facilitate reestablishment of the fungal 

assemblage relatively quickly (Jalaluddin 1969).

Studies of changes in microbial function due to fire 

are more common than direct studies of community 

structure or abundance. In western conifer systems, a 

significant proportion of the N lost during fire may be 

offset by N2 fixation by leguminous plants and their 

Rhizobium symbionts (Agee 1996). Among N-fixing 

woody species, black locust (Robinia pseudoacacia L.) 

has the most potential to contribute to the post-fire 

N economy in this way. Although black locust and its 

bacterial symbionts fix considerable amounts of N2 and 

may have been common in burned areas historically, 

this species is now most common in postagricultural old 

fields in the southern portion of the oak forest region 

(Boring and Swank 1984); thus, there are few data with 

which to assess its impact after fire.

In Georgia loblolly pine forests, considerable amounts 

of N may be fixed by herbaceous perennials (e.g., 

Desmodium spp. and Lespedeza spp.) using the same 

symbiosis (Hendricks and Boring 1999). As both of 

these genera are widespread in the Eastern United States, 

the potential exists for this pathway to be important in 

oak ecosystems. Determining whether this potential is 

achieved awaits specific studies in oak forests.

In contrast to the sparse database on N2 fixation in oak 

forests, many studies have demonstrated increases in N 

mineralization after single fires (reviews by Raison 1979; 

Boerner 1982; Wan and others 2001). These increases 

often are attributed to the alteration of organic matter by 

fire in such a manner as to render it more susceptible to 

microbial attack, to increases in microbial activity, and 

to changes in microclimate. Boerner and others (2000b, 

2004) quantified soil organic matter, N mineralization 

rate and nitrification rate in four southern Ohio mixedoak forests subjected to annual or periodic burning. 

At the full watershed scale, there was no consistent or 

persistent change in any index of microbially mediated 

N mineralization. In that study, pre-fire conditions and 

landscape characteristics were more important than fire 

behavior in explaining changes in C and N dynamics 

(Boerner and others 2000b). In a subsequent study of 

thinning+burning in a neighboring site, Boerner and 

Brinkman (2004) found no significant effect of a single 

fire on N mineralization in Ohio mixed oak-forests; a 

similar lack of fire effect on N mineralization has been 

reported for oak-pine forests in Georgia and Tennessee 

(Hubbard and others 2003) and oak-pine stands in 

North Carolina (Knoepp and others 2004).

It is important to note that the results of single fires or 

relatively short-term studies may not scale across longer 

time periods accurately. Vance and Henderson (1984) 

measured rates of N mineralization in the same long 

term Missouri stand used by Eivasi and Bayan (1996) 

a decade later. They found that N mineralization was 

reduced by longterm burning (30 years) and concluded 

that this change was a consequence of a change in 

organic matter quality rather than quantity. This is 

consistent with what one would expect if long-term 

burning results in an accumulation of black carbon over 

time (Ponomarenko and Anderson 2001).

Yet another approach to examining the impact of fire 

on the soil microbial assemblage is measuring the 

activity of the enzymes that microbes secrete into the 

soil to perform digestion processes. The activity of acid 

phosphatase is commonly used as a measure of overall 

microbial activity and as a surrogate measure for the rate 

at which N and P are mineralized from organic matter 

by microbes. Single fires in Ohio mixed-oak forests 

(Boerner and others 2000a) and pine forests in Spain 

(Saa and others 1993) and two to four fires over 5 years 

in Ohio mixed-oak forests (Boerner and Brinkman 2003) 

demonstrated reductions in acid phosphatase after fire, 

indicating an overall reduction in microbial metabolism. 

However, in the Ohio studies there were significant 

increases in phenol oxidase activity (Boerner and others 

2000a; Boerner and Brinkman 2003). Phenol oxidase 

is an enzyme produced by fungi for the breakdown of 

recalcitrant organic compounds such as those that are 

likely to be left behind after the more labile organics are 

volatilized during fire or consumed shortly after fire.
11	 Fire 4 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Again, these short-term effects may not reflect what 

might happen over a long period of fire. Eivasi 

and Bayan (1996) reported significantly lower 

microbial biomass, acid phosphatase activity, and βglucosidase activity in soils of Missouri oak-hickory 

sites exposed to annual or periodic fire for four 

decades.

Our overall understanding of the impact of fire in 

oak ecosystems on the soil microbial assemblage is 

growing but is not yet to the stage where a synthesis 

is achievable. Studies of community composition 

suggest that effects of one or several fires are rapidly 

ameliorated. Most functional studies suggest 

that the effects of fire are too modest to resolve 

analytically or are ones related primarily to changes 

in the relative proportions of labile and recalcitrant 

organic matter. However, long-term studies suggest 

that microbial activity and abundance may be 

significantly reduced if fire occurs frequently over 

many decades.

LANDSCAPE AND SCALE ISSUES

Forests with a significant oak component occur in a 

variety of landscape contexts in Eastern North America, 

from sandy coastal plains to Appalachian Piedmont and 

plateaus to the slopes of the Appalachian Mountains. 

Some of the oak region is recently glaciated, whereas 

other parts have not experienced glaciation for more 

than 106 years. Much of the eastern oak region consists 

of terrain that is dissected into ridges, hills, and valleys 

within which are strong gradients of microclimate (e.g., 

Wolfe and others 1949). In such complex landscapes, 

the behavior and effects of fire may vary so much among 

landscape positions within a watershed that considering 

only whole-watershed average effects may obscure 

important ecological patterns at the landscape scale.

To illustrate how fire behavior varies across the 

landscape, I present an example from the Ohio Hills Fire 

and Fire Surrogate (FFS) research sites in southern Ohio. 

The individual treatment units for the FFS project are 

areas of 20 to 30 ha in southern Ohio on which 50-m 

grids have been superimposed. Although the majority 

of studies that report fire behavior do so in terms of the 

maximum temperature experienced by thermocouple 

probes, this may not be the most useful metric for 

understanding fire effects. The potential impact of fire 

is better described by the integration of thermocouple 

probe temperature over the time a fire is actually present 

at a point. When expressed at a single point (above a 

single sensor) this metric is referred to as the heat index 

(Bova and Dickinson 2005) and is expressed in units 

such as oC-sec.

Figure 3 presents interpolated maps of two of the Ohio 

FFS treatment units, with a metric of landscape position 

and long-term soil moisture potential (Iverson and 

others 1997) on the left and a map of heat index on 

the right. The heat index map clearly portrays both the 

variations in fire behavior across this landscape as well as 

the strong patchiness or spatial heterogeneity typical of 

fire in these forests. Unfortunately, the time and expense 

involved in instrumenting a site for measurements 

of heat index and the unpredictability of wildfire 

occurrence combine to make measurements of actual fire 

intensity in oak forests uncommon.

Figure 3.—Interpolated maps (semivariance analysis + kriging) 

of Integrated Moisture Index (IMI) and heat index in the 

forest floor based on 50-m grid points in two southern Ohio 

watersheds. The IMI data were supplied by L. Iverson and fire 

behavior data by M. Dickinson. 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 115

The recent development of integrative, GIS derived 

metrics designed to represent within-watershed scale 

gradients of elevation, aspect, slope shape, soil type/

depth, and other aspects of microclimate have made 

available tools to allow us to begin to understand the 

landscape ecology of fire in a quantitatively rigorous 

manner. For southern Ohio, Iverson and others (1997) 

developed an Integrated Moisture Index (IMI) that 

integrates a large number of microclimate and soil 

parameters into a single metric. By stratifying sampling 

designs based on this integrated measure, we have been 

able to resolve within-watershed variations in fire effects 

on the soil. 

In Ohio mixed-oak forests where fire has been effectively 

suppressed, soil pH, inorganic N, extractable Ca2+, Mg2+, 

and Al3+, and molar Ca:Al ratio commonly vary among 

landscape positions, with relatively xeric ridgetops and 

upper, south-to-west-facing slopes having lower pH, 

cation availability, and molar Ca:Al ratio than other 

landscape positions (Table 2). N mineralization rate, 

and nitrification, and inorganic N in the soil solution 

tend to increase with increasing long-term soil moisture 

potential, or from xeric through median to mesic IMI 

classes Table 2).

By contrast, glomalin content tends to be greatest in the 

most xeric landscape positions (Table 2). As the content 

of this AM fungal-specific compound is thought to be 

directly proportional to AM fungal activity and inversely 

proportional to P availability, this suggests that the latter 

varies across the landscape in a manner parallel to that 

of N, This has been supported by direct analyses of P 

availability among landscape positions (Boerner and 

others 2003).

In some studies, relatively xeric landscape positions tend 

to have greater soil organic C content than more mesic 

positions, though others demonstrate no significant 

within-watershed scale variations in organic matter 

(Table 2). Soil enzyme activities, as indices of microbial 

activity, parallel total soil organic C closely, as do the 

abundances of various soil arthropod groups (Table 

2). Thus, variations in soil biological and chemical 

properties among landscape positions within the small 

watersheds of the Appalachian plateaus are so diverse and 

significant as to cast doubt on the utility of watershedscale averages, and point out the need for sampling 

designed to detect treatment effects to be stratified by 

landscape position.

This is illustrated by comparing responses to fire at the 

whole watershed-scale treatment unit level to those in 

the various landscape positions within that treatment 

unit (Table 3). In some cases, watershed-wide responses 

parallel (at least in direction) those in the individual 

landscape positions, e.g., phenol oxidase activity at Arch 

Rock (Boerner and others 2000a, Table 3) and both soil 

organic matter and N availability at Arch Rock (Boerner 

and others 2000b, Table 3). Far more common are 

situations in which there are significant effects of fire in 

one or more landscape positions that are not reflected 

in the overall, watershed-scale, treatment unit effect 

(Table 3), and even some where the effects at different 

landscape positions are opposite in direction, e.g., acid 

phosphatase and chitinase at Young’s Branch (Boerner 

and others 2000a, Table 3).

On level terrain with relatively small variations in 

topography and soils over fairly large areas (e.g., 

till plains and some coastal plains), extrapolating 

manipulative experimental studies on plots in the m2

 size 

range to the km2 scale at which management decisions 

are made and applied is a straightforward process of 

multiplication. By contrast, on the complex and dynamic 

landscapes of the Appalachian plateaus, such scaling up 

is more complex. GIS-based, integrative tools such as the 

IMI make the conversion from small plot-based studies 

to broad-scale prediction possible in a way that is only 

now being realized by ecologists and forest managers.
11	 Fire 6 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Table 2.—Variations in soil properties among landscape positions within Ohio mixed-oak 

forests in the absence of fire (except Boerner and others (2000b) in which fire had no significant 

effect on soil properties). The Integrated Moisture Index (IMI) classes of Iverson and others 

(1997) are used to denote landscape positions. Upper case letters reflect relative ranks, with 

A=highest and C=lowest; landscape positions with different upper case letters were significantly 

different, based on the anova or ancova used in the indicated source.

Sources/site(s) Parameter Integrated Moisture Index Class

Xeric Median Mesic

Boerner and others (2003) Soil pH B A A

Four southern Ohio Inorganic N C B A

sites pooled PO

4

3- B A A

extractable Ca2+ B A A

extractable Mg2+ B A A

extractable Al3+ A B C

Ca:Al molar ratio B A A

Morris and Boerner (1998) Soil pH B A A

Watch Rock Organic C A B B

N mineralization B B A

Nitrification B B A

Decker and others (1999) Acid phosphatase A B C

Arch Rock and Young’s Chitinase A B B

Branch pooled Phenol oxidase A A A

β-glucosidase A A A

Organic C A A A

Dress and Boerner (2001) Live root mass A A A

Arch Rock Dead root mass A A A

Root production A B B

Boerner and others (2000b) Organic C A C B

Arch Rock N mineralization C B A

Nitrification B B A

Boerner and others (2000b) Organic C A A A

Young’s Branch N mineralization C B A

Nitrification B A A

Knorr and others (2003) Glomalin content C B A

Arch Rock

Dress and Boerner (2004) Oribatid Mites A B B

Arch Rock Collembola A A A
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 117

CONCLUSIONS

The last century has seen literally thousands of individual 

studies of the effects of fire on the forest floor and soils 

of American forests, shrublands, and grasslands. Despite 

this rich literature, the available database for the eastern 

oak forest ecosystems is surprisingly sparse, particularly 

in studies that have continued past the first post-fire year. 

Most fires in oak ecosystems for which there are data 

generally have intensities that fall in the lower end of 

the range for North American ecosystem fires, though 

occasional higher intensity, stand-replacing fires have 

been reported. Most of the unconsolidated litter and a 

portion of the humus layer are combusted during typical 

oak forest fires, yet significant heating of the mineral soil 

is limited to areas where accumulations of fuel smolder 

for extended periods. As a result, direct mortality of 

organisms in the soil is low, and mortality in the forest 

floor is proportional to the extent of consumption.

A considerable portion of the N in detritus and 

vegetation may be lost to a combination of direct 

volatilization and ash convection. Ash convection also 

can result in significant losses of P and base cations. 

Fire behavior, intensity, and weather are important 

regulators of these processes. The nutrients contained 

in ash typically are dissolved by the first several 

precipitation events following fire, and the proportion 

of those solubilized nutrients that remain in the soil for 

subsequent plant use depends on the exchange capacity 

of the soil, landscape geomorphology, and weather. 

In shrublands and coniferous forests, development of 

hydrophobicity in soil during and just after fire is often 

an important factor in determining how much of the 

post-fire precipitation and solubilized nutrient load are 

lost to runoff. However, the literature does not suggest 

that this is an important process in eastern oak forests.

The availability of N, a potentially limiting nutrient 

for post-fire plant growth, and the total amount of 

organic matter in the soil are typically greater after fire, 

though this may be a more transient phenomenon in 

oak forests than in coniferous forests and shrublands 

where fire intensities are considerably greater. The 

Table 3.—Fire-induced variations in soil properties at the watershed scale and among landscape 

positions within Ohio mixed-oak forests. The Integrated Moisture Index (IMI) classes of Iverson 

and others (1997) are used to denote landscape positions, and the column labeled “Pooled” 

represents the full watershed. Effects of fire are indicated as 0=no significant effect, ↓=significant 

decrease, and ↑=significant increase, based on the anova or ancova used in the indicated source.

Sources/site(s) Parameter Integrated Moisture Index Class

Pooled Xeric Median Mesic

Boerner and others (2000b) Organic C 0 0 0 0

Arch Rock Total Inorganic N 0 0 0 0

Boerner and others (2000b) Organic C 0 ↑ ↑ 0

Young’s Branch Total Inorganic N 0 0 ↑ 0

Boerner and others (2000a) Acid phosphatase ↓ 0 ↓ 0

Arch Rock Chitinase 0 0 0 0

Phenol oxidase ↑ ↑ ↑ ↑

Boerner and others (2000a) Acid phosphatase ↓ ↓ ↓ ↑

Young’s Branch Chitinase ↓ ↓ 0 ↑

Phenol oxidase ↑ 0 0 ↑

Dress and Boerner (2001) April Root Mass ↓ 0 0 ↓

Arch Rock May Root Mass 0 0 ↑ 0

June Root Mass ↓ 0 0 ↓
11	 Fire 8 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

possible importance of the charcoal (black carbon) 

remaining after partial combustion of woody material 

to post-fire biological activity is anl area of current fire 

research in coniferous forests and grasslands, though little 

information on black carbon in oak forests is available.

Microbial activity, biomass, and community structure 

in the forest floor often is affected in proportion to the 

consumption of this organic-rich layer. By contrast, 

short-term effects on microbes in the mineral soil seem 

to be primarily the result of progressive soil heating 

under accumulations of organic materials that smolder 

for longer periods. Recovery of the microbial assemblage 

from fire is relatively rapid and is the result both of 

propagules in situ and colonization from surrounding, 

unburned areas. Similar spatial and temporal patterns 

have been reported for microarthropods in the forest 

floor and soil, but the database on other soil invertebrates 

groups is so sparse as to warrant no generalization.

Much of the oak forest region of Eastern North America 

is complex in geomorphology, and this complexity is a 

challenge when experimental studies must be scaled up 

to land areas as large as those for which management 

decisions are made. The use of GIS-derived measures of 

landscape structure and heterogeneity to stratify sampling 

designs, inform statistical analyses, and integrate plot 

results to the landscape scale has emerged as a viable tool 

for generating management-scale information.

ACKNOWLEDGMENT

This review could not have been possible without the 

many years of collaboration, discussion, argument, and 

debate in which I’ve been privileged to participate. Elaine 

Kennedy Sutherland, Todd Hutchinson, Louis Iverson, 

Dan Yaussy, Jennifer Brinkman, Bill Dress, Sherri Morris, 

and Tom Waldrop have been particularly important in 

this regard. Betsy Wrobel-Boerner, Matthew Dickinson, 

and two anonymous reviewers improved the quality of 

this manuscript significantly. This is publication No. 80 

of the Fire and Fire Surrogate Network Program. The 

financial support of the USDA Forest Service’s Ecosystem 

Management Program, the U.S. Joint Fire Science 

Program, National Fire Plan, and National Science 

Foundation is gratefully acknowledged. 

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Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 123

Responses of Oak and Other Hardwood Regeneration to

Prescribed Fire: What We Know as of 2005

Patrick H. Brose1†, Thomas M. Schuler2, and Jeffrey S. Ward3

Abstract.—An obstacle to using prescribed fire to manage mixed oak forests is the varied results of previous 

fire studies. It has been reported that fires enhanced, hindered, or had no effect on the competitive position of 

oak in the regeneration pool. We review a portion of the published literature and identify key factors that led 

to the relative competitiveness of oak reproduction benefiting from or being harmed by prescribed fires. These 

key factors are synthesized into general guidelines to help practitioners understand how fire can be a positive 

force in the oak regeneration process. We also point out some situations where fire hinders the competitive 

position of the oak regeneration, and provide suggestions for researchers studying fire in mixed oak forests.

fire intensities, associated silvicultural practices, and 

number of fires, and include an equally wide range of 

results. The competitive position of oak reproduction in 

the regeneration pool can be enhanced or hindered or 

not be affected. In this paper we review a portion of the 

fire/oak literature, grouping studies during the past four 

decades by stand structure (fully stocked mature and less 

than fully stocked mature), fire intensity (low and high), 

number of burns (single, twice, three to five, and six or 

more), and response of oak regeneration. We synthesize 

the common denominators for when the competitive 

position of oak reproduction is enhanced or harmed by 

prescribed fire and provide guidelines on using fire to 

help regenerate mixed oak forests.

Single Fires

Mature Stand, Low-Intensity Fire

This is by far the most common stand and fire type 

in the literature (Fig. 1). In this group, mature stands 

are in the understory reinitiation stage of development 

(Oliver and Larson 1996), usually are at least 75 years 

old and are near the end of a typical timber rotation 

for oak sawlogs. Low-intensity fires are defined as those 

with flames less than 2-feet long, which consume only 

the leaf litter and small woody debris, and cause little, 

if any, overstory mortality. The effect of these fires in 

this setting may be beneficial, detrimental, or neutral 

with respect to the relative competitiveness of oak 

regeneration (Fig. 1). American beech (Fagus grandifolia) 

remained the dominant species after a light surface fire 

in New York (Swan 1970). Nyland and others (1982) 

reported that the dense understory that develops after a 

single fire in New York could inhibit oak regeneration. 

Introduction

There is developing consensus that fire has had a close 

association with and has shaped the species composition 

of mixed-oak forests in the eastern United States for the 

last several thousand years (Crow 1988; Abrams 1992; 

Brose and others 2001). Scars on stem cross sections are 

evidence that prior to the fire control era that began in 

the 1920s, fires in Ohio, Maryland, and West Virginia 

occurred at intervals of 5 to 15 years during the previous 

four centuries (Sutherland 1997; Shumway and others 

2001; Schuler and McClain 2003). Other fire-scar 

studies have documented fire return intervals ranging 

from 2 to 24 years from New Jersey to the western 

portion of the mixed-oak forest in Missouri (Buell 1953; 

Dey and others 2004). Evidence of reoccurring fire 

also predates European contact in the eastern United 

States. In a charcoal and pollen accumulation study 

in the southern Appalachians, Delcourt and Delcourt 

(1997) found evidence of landscape-level fires in a 

forest dominated by oak, American chestnut, and pine 

throughout the past 3900 years.

The heretofore unrecognized close association of fire 

and oak forests in the eastern United States has led 

researchers to investigate the use of prescribed fire to 

sustain and regenerate oak forests. Eastern fire studies 

cover a wide range of stand characteristics, species mixes, 

1

USDA Forest Service, Northeastern Research Station, Irvine, 

PA 16329; 2USDA Forest Service, Northeastern Research 

Station, Parsons, WV 26287; 3Connecticut Agricultural 

Experiment Station, Department of Forestry and Horticulture, 

New Haven, CT 06504; †Corresponding author, 814-563-1040, 

email: pbrose@fs.fed.us.
12	 Fire 4 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

In a study of the same burns, McGee and others 

(1995) found that the single fire increased 

the competitive status of American beech and 

red maple (Acer rubrum) while reducing the 

importance value of northern red oak (Quercus 

rubra). Prescribed fire reduced northern red 

oak survival and did not increase basal diameter 

growth in northern Georgia (Loftis 1990a). 

Low-intensity fires increased red maple and black 

birch (Betula lenta) density relative to oak in 

Connecticut (Ducey and others 1996). In this 

study there was no difference in height growth 

of oaks between the low-intensity and unburned 

sections (Moser and others 1996).

One of the more widely reported studies is the 

National Fire and Fire Surrogates project in 

southern Ohio. Low-intensity fires in unmanaged 

stands did not increase the number of oak 

seedlings after 1 year (Iverson and others 2004) or 2

years (Apsley and McCarthy 2004). In an associated 

study, McQuattie and others (2004) found that burning 

had no significant effect on seedling height of black 

oak (Q. velutina) and red maple. However, in this 

study, seedlings of both species were more massive on 

burned than unburned plots. A later report of the same 

study found that red maple seedlings, but not those of 

chestnut oak (Q. prinus), were taller in burned than 

unburned sections 2 years after the burn (Apsley and 

McCarthy 2004). In a related study also in southern 

Ohio, Hutchinson and Sutherland (2000) reported 

that red maple and white oak seedling frequencies were 

unchanged 3 years after a light surface fire. The effect 

of low-intensity fire on comparative height growth was 

equally ambiguous.

Several studies of low-intensity fire in mature stands 

have reported beneficial effects on oak regeneration. 

Barnes and Van Lear (1998) found that 2 years after a 

low-intensity spring fire in the southern Appalachians, 

regeneration (less than 2.0 inches in d.b.h.) densities of 

oak, hickory (Carya spp.), and sourwood (Oxydendrum 

arboreum) increased, while red maple density decreased. 

This change in species composition resulted in a slight 

increase in relative density of oak regeneration from 31

percent before the prescribed burn to 37 percent 2 years 

later. In Kentucky, Adams and Rieske (2001) found 

that the seedling height and diameter growth of white 

oak (Q. alba) was enhanced by a prescribed burn. A 

separate study on Kentucky ridgetops reported increases 

in density of both oak and red maple seedling densities 

after prescribed fire (Kuddes-Fischer and Arthur 2002). 

A low-intensity fire in south-central New Jersey resulted 

in low mortality of small oaks (Little 1946).

Mature Stand, High-Intensity Fire

In this category, high-intensity fires are defined as those 

with flames longer than 2 feet or killing significant 

numbers of overstory trees. Studies reporting on this type 

of fire in mature stands are less common than for the 

previous group (Fig. 1). Generally, these fires enhanced 

the competitive status of oak regeneration. Several of 

the researchers examined the aftereffect of wildfires by 

comparing burned areas and adjacent unburned stands. 

In New York, high-intensity fires did not change the 

average frequency of black oak but reduced the average 

frequency of red maple and white oak (Swan 1970). 

Swan also reported that the percentage of oaks top-killed 

by the fire was lower than that for red maple, sugar 

maple (Acer saccharum), and American beech.

Both absolute and relative densities of oak were increased 

7 years after a high-intensity prescribed fire in North 

0

20

40

60

80

100

120

Mature - Low Mature - High Cut - Low Cut - High

Stand Structure - Fire Intensity

Reviewed Studies (%)

Benefits oak Neutral Harms oak

14 Studies 10 Studies 7 Studies 5 Studies

Figure 1.—Distribution of reviewed studies of the response of 

hardwood regeneration to single fires. The studies were divided into 

12 groups based on stand structure, i.e., had the overstory been 

harvested to some degree, reported fire intensity (flame length or 

overstory mortality), and whether the fire benefited, harmed, or did 

not affect the competitive position of oak in the regeneration pool.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 125

Carolina (Maslen 1989). Although survival of oaks 

within a given size class was higher than for non-oak 

species following the fire, height growth was greater for 

non-oak than for oak species. Another North Carolina 

study reported that the density of scarlet (Quercus 

coccinea) and chestnut oak increased 1 year after a highintensity fire while the density of red maple, northern 

red oak, and white oak decreased (Elliott and others 

1999). In Connecticut, Ducey and others (1996) 

reported the density of northern red oak, eastern white 

pine (Pinus strobus), and black birch was higher 7 years 

after a high-intensity prescribed burn than on unburned 

plots while the density of red maple and white oak 

was decreased by the fire. As a follow-up to the Ducey 

paper, Moser and others (1996) reported that the height 

growth of oaks was greater following the high-intensity 

fire than that of oaks in the unburned controls.

Perhaps the primary reason for the increase in survival 

and growth of seedlings after high-intensity fires is that 

the amount of light reaching regeneration following fireinduced mortality of overstory trees was increased. Loftis 

(1990b) reported that the survival of northern red oak 

seedlings increased following a reduction in overstory 

density (similar to a high-intensity fire) as opposed to 

a reduction in understory density (similar to a lowintensity fire).

High-intensity fires are not a panacea for regenerating 

oaks. In addition to the resulting economic loss of 

timber (Hepting 1941; Little 1946; Paulsell 1957; 

Loomis 1973; Ward and Stephens 1989), high-intensity 

fires may inhibit the development of oak regeneration 

relative to other species. In West Virginia, the density of 

northern red and chestnut oak advanced regeneration 

decreased 5 years after a fire that reduced overstory 

basal area by 17 percent (Wendel and Smith 1986). In 

contrast, red maple density increased slightly over the 

same period.

Cut Stand, Low Intensity Fire

This stand type differs from the mature type in that 

cutting by group selection, shelterwood, clearcut, 

etc preceded the fire by one or more years and the 

regeneration layer was developing into saplings, i.e., the 

stand initiation stage (Oliver and Larson 1996). There 

were fewer studies of fire for this stand than for mature 

stands (Fig. 1). 

In West Virginia, Collins and Carson (2003) reported 

that prescribed burning of recently created gaps reduced 

the abundance of northern red oak while the density of 

competitors was increased or not affected. Dolan and 

Parker (2004) found that a combination of burning 

and thinning did not result in the establishment of new 

oak seedlings on mesic or xeric oak sites in Indiana, 

but did result in greater consistency with respect to the 

number of shade-tolerant and intolerant seedlings in 

the understory. Jackson and Buckley (2004) noted an 

increase in small oak seedlings but attributed this to the 

sprouting of large oak seedlings that had been top-killed 

by fire a year earlier.

Fire studies in cut stands with a longer interval between 

the harvest and fire have shown substantial benefits 

to the oak regeneration. Brose and Van Lear (1998) 

and Brose and others (1999a) reported increases in 

the relative dominance of oak when burning was done 

4 years after a heavy shelterwood cut. Kruger and 

Reich (1997) reported comparable results from fires 

conducted in 4-year-old oak shelterwoods in Wisconsin. 

In Connecticut, burning several years after a heavy 

shelterwood cut or a complete overstory removal also 

has increased the competitive position of oak in the 

regeneration pool (Ward and Gluck 1999; Ward and 

Brose 2004).

Cut Stand, High Intensity Fire

This grouping of papers was the smallest (Fig. 1) but 

included the oldest fire study in the eastern United 

States. A high-intensity fire in 1932 topkilled half of the 

stems less than 6 inches d.b.h. in a 30-year-old sapling 

stand in Connecticut. Fifty-five years after that fire, Ward 

and Stephens (1989) reported that oak densities were 

twice as high in burned than in unburned areas. The fire 

had no long-term effect on the density of other species 

groups. Comparable results were reported for Rhode 

Island (Brown 1960).

One of the first attempts to use fire along with a 

partial reduction in overstory stocking to increase the 

importance of oak in the understory was by Wendel 
12	 Fire 6 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

and Smith (1986) in the Ridge and Valley 

region of eastern West Virginia. The effects 

of the fire were evaluated on the overstory 

and understory. Two years before the fire, 

the stand was thinned from below, e.g., the 

undesirable trees less than 6 inches d.b.h. plus 

cull trees were felled and left on the site. Strip 

head fires in the spring of 1980 consumed 

about 56 percent of the litter fuels and 18 

percent of the woody fuels. Fire intensity 

varied with strip width, fuel loading, and wind 

patterns, though it likely was increased by 

the addition of fuels from the thinning 2 years 

earlier. Average fire scar height was about 5 feet, 

indicating a high-intensity fire for hardwood 

leaf litter. The fire reduced the number of stems 

more than 5 inches d.b.h. from 115 to 93 per 

acre and reduced basal area from 90 to 73 feet/acre. In 

the understory, red maple and black locust seedlings 

and sprouts increased in number while the number of 

northern red and chestnut oak decreased.

In a fire study conducted in the Virginia Piedmont, 

Brose and Van Lear (1998) and Brose and others (1999a) 

found that high-intensity fire was especially beneficial 

to the relative competitiveness of oak regeneration if the 

fire occurred during the spring versus summer or winter. 

High-fire intensity also was more beneficial to oak than 

low-intensity fire in studies by Ward and Gluck (1999) 

and Ward and Brose (2004).

Multiple Fires

Multiple prescribed burns at interval of 2 to 3 years were 

suggested by Sander (1988) to control unwanted woody 

competition. However, the impact of multiple burns 

may vary by the number of burns and their frequency 

(Fig. 2). Some researchers reported that periodic burns 

increase the competitive status of oak regeneration. A 

prescribed fire in Kentucky followed 2 years later by 

a wildfire increased the number of oaks in the shrub 

stratum (20 inches tall to 1 inch d.b.h.) more than six 

fold (Arthur and others 1998). This combination of 

fires also doubled the number of red maple in the shrub 

stratum versus the unburned plots. However, the plots 

that had been burned only once had higher red maple 

densities than the plots that had been burned twice. The 

second fire reduced the density of red maple sprouts and 

increased the number of chestnut oak sprouts.

In New York, both the relative and absolute densities of 

northern red oak reproduction more than 4.5 feet were 

greater on plots that had burned twice than on plots 

that had been burned once in New York (McGee and 

others 1995). This study also reported that northern red 

oak densities were higher following two high-intensity 

than two low-intensity fires and that the two hot burns 

increased the competitive status of American beech. 

Periodic burns at intervals of 5 years or less have nearly 

eliminated red maple from red pine (Pinus resinosa) 

stands in Michigan (Henning and Dickmann 1996).

Barnes and Van Lear (1998) reported that after three 

winter burns in South Carolina, there were fewer oaks 

than after a single spring fire. However, the winterburned stands contained more than 5,000 stems/acre of 

oak regeneration and the height of oak sprouts following 

three burns was more than twice that of sprouts after a 

single spring burn. Following three periodic prescribed 

fires in Tennessee, oak regeneration was abundant, 

averaging more than 1,500 stems/acre (Clatterbuck 

1998). Basal diameter and relative height growth of 

white oak seedlings were increased by two prescribed 

burns in Kentucky (Adams and Rieske 2001). However, 

relative height growth was greatest for seedlings in plots 

burned only once.

0

20

40

60

80

100

120

2 Burns 3-5 burns 6+ Burns Annual Burn

Reviewed Studies (%)

Benefits oak Neutral Harms oak

6 Studies 3 Studies 5 Studies 3 Studies

Figure 2.—Distribution of reviewed studies of the response of 

hardwood regeneration to multiple fires. The studies were divided into 

12 groups based on the number of periodic burns and whether the fire 

benefited, harmed, or did not affect the competitive position of oak in 

the regeneration pool.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 127

Other researchers maintain that periodic burning should 

be used with caution if the management objective is 

to regenerate oak. In a study in Tennessee, Thor and 

Nichols (1973) found that two periodic burns separated 

by 5 years increased the density of oak, sumac (Rhus

spp.), and sassafras (Sassafras albidum) regeneration and 

eliminated red maple reproduction. However on the 

same plots, four additional periodic burns at intervals 

of 4 to 5 years the same plots increased the absolute 

and relative densities of red maple (DeSelm and others 

1991). Concurrently, the density of post (Q. stellata), 

scarlet, and southern red oak (Q. falcata) decreased. 

In Missouri, periodic burning reduced the density of 

small oak saplings (Sasseen and Muzika 2004) while 

oak mortality increased with burning frequency (Dey 

and Hartmann 2004). For example, a single fire only 

resulted in 10-percent mortality among black oaks with 

a basal diameter of 1 inch, but three or more fires killed 

25 percent of stems. In New Jersey, the mortality of 

oak saplings (one inch d.b.h) increased as the intervals 

between fires decreased. Mortality was 100 percent 

on annually burned plots, 62 percent on plots burned 

at 3-year intervals, and 11 percent mortality on plots 

burned at intervals of 10 years or longer (Somes and 

Moorhead 1950). In Kentucky, Green and others (2004) 

reported that red maple seedlings were tallest on plots 

burned three times than on those burned twice or not 

at all. They also noted that oak seedlings were tallest on 

unburned plots. Although red maple seedlings were taller 

than oak seedlings, mortality was higher in red maple.

In most studies researchers reported that annual burning 

reduces all regeneration density, including oak seedlings 

and saplings (Little and Moore 1949; Chaiken 1952; 

Paulsell 1957; Grano 1970). Although 6 years of annual 

burning in Tennessee increased total regeneration density 

and the density of oak regeneration (Thor and Nichols 

1973), an additional 21 years of annual burns on these 

same plots eliminated all regeneration except for winged 

sumac (R. copallina) and oak (DeSelm and others 1991). 

Although some oak remain (primarily post oak) total 

density has declined by 97 percent or more. In the 

Missouri Ozarks, four annual burns greatly reduced or, 

in one case, eliminated small oak saplings less than 1.5 

inches d.b.h. (Sasseen and Muzika 2004) and 14 years 

of annual burning eliminated large sapling stems in 

Minnesota (White 1983). The frequency of red maple 

seedling decreased after three annual burns in southern 

Ohio (Hutchinson and Sutherland 2000); surprisingly 

the frequency of white oak seedlings was not changed by 

the annual burns.

Synthesis

Every possible response of oak regeneration to fire is 

found in the literature, though commonalities among 

studies are apparent when the competitive position of 

the oak reproduction is enhanced or hindered. Results 

were most varied for single, low-intensity prescribed fires 

in mature stands. Where oak regeneration was harmed 

in this setting (Loftis 1990a; Nyland 1982; McGee and 

others 1995; Moser and others 1996), the studies were 

conducted on high-quality sites. The oak stems were 

small and they had been growing in dense shade. These 

stems probably had small root systems and minimal root 

carbohydrate reserves, and likely were of low vigor at the 

time of the fire. Several researchers have reported that 

such stems, regardless of species, usually do not survive 

fire of any intensity (Johnson 1974; Loftis 1990a; Brose 

and Van Lear 2004). Also, the fires did not significantly 

increase the amount of sunlight reaching the forest floor, 

so oak stems sprouting after the fires were forced to grow 

new tissue in sub optimal light conditions.

When single low-intensity surface fires benefited oak 

regeneration in mature stands (Little 1946; Barnes and 

Van Lear 1998; Adams and Rieske 2001; Kuddes-Fischer 

and Arthur 2002), the studies were conducted on dry, 

hot, sandy soils or xeric upper slopes. Oak regeneration 

tends to accumulate on such sites because they have 

sparser canopy cover and more sunlight reaches the forest 

floor (Johnson and others 2002). Thus, the regeneration 

has larger root systems and greater root carbohydrate 

reserves, and is more vigorous. Brose and Van Lear 

(2004) found that such oak regeneration generally 

sprouts after a low-intensity fire.

Single low-intensity fires in cut stands also produced 

varied results. The major differences between studies 

reporting a positive effect on oak regeneration and those 

reporting no effect or a negative impact were related to 

the intensity of cutting and the length of time between 

the cutting and the fire. The positive studies (Kruger 
12	 Fire 8 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

and Reich 1997; Brose and Van Lear 1998; Ward and 

Gluck 1999; Ward and Brose 2004) were conducted in 

heavily cut shelterwoods or clearcuts and at least 4 years 

had passed between the cutting and the fires. In the 

neutral and negative studies (Collins and Carson 2003; 

Dolan and Parker 2004; Jackson and Buckley 2004), the 

degree of cutting was much lighter e.g., small gaps and 

low thinning, and only 1 or 2 years elapsed between the 

cutting and the fire. There is a clear relationship among 

the amount of light, number of growing seasons, and 

size/vigor of oak seedlings (Miller and others 2004). In 

the positive studies, the oak reproduction at the time of 

the cutting had sufficient light and time to build large 

root systems so it was more likely to sprout after the fires. 

In the neutral and negative studies, the regeneration 

reacted like small, low-vigor seedlings experiencing fire in 

a mature stand: it died or responded poorly.

The high-intensity fires in mature and cut stands 

generally benefited oak by being more detrimental to 

the survival and growth of non-oak species than to the 

oaks. This disparity is attributed to inherent differences 

in germination strategy between oaks and many of their 

competitors. Acorns have hypogeal germination, i.e., 

cotyledons remain in the shell and serve as a belowground energy source for seedling development. Seeds 

of common competitors, e.g., red maple, have epigeal 

germination, i.e., cotyledons emerge and rise above 

the shell to form the first photosynthetic leaves. This 

difference in germination strategy places oak seedlings’ 

root collar, and the accompanying dormant buds, lower 

than that of red maple. This difference in germination 

strategy is accentuated by wildlife. Acorns are routinely 

buried an inch or more into the forest floor by birds 

and small mammals while seeds from red maple and 

other competitors typically are not cached. Thus, an 

oak seedling generally has a deeper root collar than a 

red maple seedling due to seed burial and hypogeal 

germination. Consequently, hotter fires are more likely 

to kill the dormant buds of red maple than those of oak.

Another important silvical difference between oak 

regeneration and the reproduction of many competitors 

is the developmental rate of the root system. Upon 

germinating, oaks send a strong radicle deep into the soil 

to establish a taproot and emphasize root development 

over stem growth (Kelty 1988; Kolb and others 1990). 

Most competitors take the opposite approach; root 

development is sacrificed to promote rapid stem growth. 

Thus, oak regeneration usually has shorter stems than 

those of the competition but has larger root systems. 

Because of these silvical characteristics – hypogeal 

germination, emphasis on root development, and seed 

burial by wildlife – oak regeneration is less detrimentally 

affected by high intensity fires than its competitors.

 The adverse effect of high-intensity fire on oak 

regeneration in previously cut stands (Wendel and Smith 

1986) likely was caused by the short interval between 

cutting and the fire. Also, a high deer population may 

have preferentially browsed oak sprouts more than those 

of competitors during the 5 years between the fires and 

data collection.

Generally, two to five fires spread over a decade or 

more have benefited the competitive position of oak 

in the regeneration pool, likely due to selecting against 

species with inherently smaller root systems and 

creating/maintaining an understory light regime more 

favorable to the development of oak rootstocks. Multiple 

periodic burns may be a reasonable mimic of single, 

high-intensity fires in that they both create more open 

understories (but at different time scales). 

However, multiple periodic fires do not always benefit 

oak. Merritt and Pope (1991) compared single burns 

conducted in the fall with a second burn conducted in 

the spring two growing seasons later. Consistent with 

other studies, both the first and second prescribed fires 

significantly reduced the number of competing shadetolerant saplings 10 to 20 feet tall; however, neither the 

first or second prescribed burn aided in the establishment 

of new oak seedlings. Merritt and Pope concluded that 

it was not advisable to use fire alone, including multiple 

fires, to aid in the establishment of new oak seedlings in 

stands described as “recently mature.”

Although the root mass/size of oak regeneration 

dwarfs that of most competitors of similar-size, it can 

be exhausted by multiple resprouting. This is evident 

in studies in which multiple periodic fires or annual 

burning were examined. There seems to be a point at 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 129

which repeated fire does not continue to benefit the oak 

component of the regeneration pool. Where that point 

is and how rapidly it is reached likely is a function of 

several stand characteristics.

None of the reviewed fire/oak papers focused exclusively 

on oak and all contained information on other eastern 

species. From these papers, species can be grouped 

into three broad classes: susceptible, intermediate, and 

resistant to fire. Susceptible species, which are poor 

sprouters following fire, include black birch, eastern 

hemlock (Tsuga canadensis), eastern white pine (P. 

strobus), hophornbeam (Ostrya virginiana), sugar maple, 

and yellow-poplar (Marquis 1975; Kruger and Reich 

1997; Brose and Van Lear 1998; Ward and Brose 2004). 

Fire-resistant species include the oaks, sympatric species 

such as the hickories (Carya spp), and those capable of 

consistent root sprouting e.g., aspen (Populus spp.), black 

locust (Robinia pseudoacacia), and sassafras, as long as 

there is sufficient light for sprouts to survive (Kruger 

and Reich 1997; Brose and Van Lear 1998; Waldrop 

and Brose 1999). Intermediate species differ from 

resistant species in how their sprouting ability responds 

to changes in fire intensity, fire seasonality, and seedling 

size. This is particularly true of red maple (Huddle and 

Pallardy 1996, 1999). Small red maple is readily killed 

by fire, even low intensity burns (Reich and others 1990) 

but the sprouting ability of this species develops as size 

increases (Brose and Van Lear 2004; Ward and Brose 

2004). In most dormant-season burns and low-intensity 

fires, large advance red maple regeneration equals that 

of oaks in sprouting ability (Brose and Van Lear 1998). 

However, red maple is a markedly poor sprouter as fire 

intensity increases, especially during the growing season. 

Other intermediates that exhibit a wide range of root 

sprouting ability include American beech, blackgum 

(Nyssa sylvatica), and pin cherry (Prunus pensylvanica).

Guidelines for Foresters and 

Researchers

The following guidelines can aid foresters who plan to 

use fire to regenerate oak forests: 

1. Prescribed fire is best used after significant 

reductions in overstory stocking to release 

existing oak seedlings, sometimes referred to 

as advanced regeneration, from competition. 

Prescribed fires should occur after released oak 

seedlings have had time to develop robust root 

systems (root collar diameter > 0.75 inch) that 

can sprout vigorously. The recommended time 

is 3 to 5 years after a shelterwood harvest that 

reduces stocking by about 50 percent in a fully 

stocked stand (Brose and others 1999a, b). 

Measures to protect valuable residual trees likely 

will be needed (Brose and Van Lear 1999).

2. Prescribed fires in late spring or summer are 

the most lethal to oak competitors because their 

root reserves are at their lowest levels. In some 

locations, the window of opportunity for such 

burns may be too short for planning purposes, 

so the focus should be on spring burns just 

before or during leaf expansion.

3. In the absence of adequate numbers of existing 

oak seedlings, the use of fire to establish new oak 

seedlings generally has not been successful. Some 

factors related to understories with inadequate 

densities of advanced oak regeneration are not 

related to the fire and oak relationship. Fire can 

create conditions suitable for oak establishment 

by reducing litter layers and reducing 

understory and midstory competition, but in 

the absence of large acorn crops, the newly 

available growing space will be used by other 

species in 1 to 2 years. Seedling establishment 

is largely dependent on large acorn crops that 

can overwhelm seed and seedling consumers, 

for example insects, rodents, and white-tailed 

deer. Avoid burning when an acorn crop has 

just occurred (Auchmoody and Smith 1993) or 

when oak seedlings are small, of low vigor, or 

recently established (Loftis 1990a; Brose and 

Van Lear 2003). 

4. The use of fire following clearcuts has produced 

encouraging results in some locations. When 

oaks are present in the regeneration but 

other species are at much higher densities, a 

moderately intense prescribed fire can increase 

the relative density of oak by taking advantage 
1	 Fire 30 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

of this species’ superior sprouting characteristics. 

Like shelterwood harvests, prescribed fires after 

clearcuts should be scheduled to allow oaks 

time to develop robust root systems that can 

produce vigorous sprouts. The time needed 

may be less than the 3 to 5 years recommended 

for the shelterwood regeneration process, but 

allow sufficient time for seeds of other species 

to germinate. Fires in clearcuts do not injure 

valuable residual trees unless some are retained 

for other management objectives. 

5. Deer browsing is important in many areas 

and must be addressed should densities exceed 

a critical threshold before the regeneration 

process. High deer densities drive forest 

succession and discriminate against sprouting 

oaks after a fire (Collins and Carson 2003; 

Horsley and others 2003). This is an added 

obstacle on productive mesic sites where 

competition from non-oaks is greatest. In such 

cases, the use of fencing or other drastic means is 

necessary to mitigate the impact of deer. 

6. A successful oak regeneration plan likely will 

ensure a diversity of woody species in the new 

stand. None of the silvicultural studies we 

reviewed reported the elimination of common 

oak competitors though many documented 

the virtual loss of oak when the regeneration 

plan favors other species intentionally or 

unintentionally.

The following are reminders concerning the essentials 

of high-quality research as well as points to consider in 

conducting fire research in oak forests:

1. Classify or measure fire intensity and/or severity 

at the same scale used to monitor regeneration 

response. This was by far the most common 

shortcoming in the studies we reviewed. The 

authors of several early fire/oak papers did not 

describe fire behavior while others included 

only general descriptions (cool, hot) at the stand 

level. Fortunately in more recent papers, fire 

behavior is described in more appropriate terms, 

for example, flame length, rate of spread, and 

char height, to help the reader gain a mental 

picture of the fire. The best papers report fire 

behavior and fuel consumption at the same scale 

that responses are measured. 

2. Document preburn forest conditions. The 

response of a plant community to fire is 

influenced by several preexisting factors, 

especially the size/vigor of the regeneration and 

understory light levels. The more the preburn 

conditions are documented, the more useful the 

study. For oak reproduction this almost always 

requires measuring the diameter of the root 

collar

3. Use a control treatment. This should seem 

obvious, but, we reviewed studies that did 

not include unburned controls. Statistical 

inference, subsequent publication, and value to 

the scientific community of fire studies will be 

greatly enhanced with a control.

4. Do not be hasty in reporting results. Fire 

effects may take three years or more to be fully 

manifested. Early results need to be reported as 

preliminary rather than as definitive.

5. Publication of fire/oak studies need to be in 

high-quality journals so they reach the scientific 

community and technical outlets for the 

practicing forester.

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1	 Fire 36 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

FIRE AND THE HERBACEOUS LAYER OF EASTERN OAK FORESTS

Todd Hutchinson1

Abstract.—Across oak forest landscapes, the herbaceous layer supports the great majority of plant 

diversity. As the use of prescribed fire increases, it is important to better understand its effects on 

biodiversity. This paper reviews the current “state of the knowledge” regarding fire effects on herbaceous 

layer vegetation. In typical dormant-season fires, direct heating effects are minimal on most herbaceous 

plants (forbs, grasses, sedges). Although woody plants are topkilled, nearly all resprout. Fire indirectly 

affects the herb layer by altering the forest floor and soil environments. The consumption of leaf 

litter during fire stimulates germination for a number of seedbanking species. Three case studies (oak 

forests in Missouri and Ohio, oak barrens in Illinois) of herb-layer response to fire are reviewed. These 

and other studies show that species richness and the cover of herbaceous plants usually increase after 

fire. Fire can have unique effects on herbaceous communities that are not realized with mechanical 

treatments (e.g., partial harvesting) alone. Although prescribed fire is commonly applied to maintain 

open-structured habitats that often contain rare plants, it also could be a useful management tool 

for sustaining and enhancing rare plant populations in upland oak forests. What is lacking most 

from our knowledge of how fire regimes affect the herbaceous layer of oak forests is: 1) the longterm effects of fire suppression, and 2) the long-term effects of periodic application of prescribed fire.

most rare plant species. The herb layer also provides 

habitat and food for numerous species ranging from 

arthropods to large mammals.

Broadly defined, herbaceous life forms are forbs (broadleaved plants) and the graminoids (grasses and sedges); 

woody life forms are trees (seedlings and sprouts) 

and shrubs (including woody vines). Perennial forbs 

comprise the majority of plant diversity: in Ohio oak 

forests, perennial forbs made up 60 percent of species, 

followed by graminoids (14 percent), trees (11), shrubs 

and woody vines (9) and annual forbs (6) (Hutchinson 

et al. 2005a). However, woody plants, often have greater 

cover and biomass than herbaceous plants, particularly 

on dry sites (e.g., Hartman and Heuman 2003). Another 

characteristic of herb-layer vegetation is that most of the 

species occur infrequently across the landscape (Keddy 

2005).

Many factors affect the composition and diversity of 

herb-layer communities in oak forest landscapes. At the 

broad scale, plant assemblages vary with climate and 

landforms while at the local scale (e.g., a watershed), 

species abundances are strongly associated with 

topographic gradients of soil moisture and fertility 

(Hutchinson et al. 1999). Also within oak forest 

landscapes, areas where tree growth is restricted (e.g., 

INTRODUCTION

The objective of this paper is to review what is known 

about the response of herbaceous layer vegetation to 

fire in eastern oak landscapes. While the focus is on 

fire effects in oak forests in the central hardwoods 

region, more open canopied communities within oak 

forest landscapes are included, as is research from other 

ecosystems, particularly when information is lacking 

from oak forests. In reviewing the literature, two major 

generalizations became evident: 1) although there have 

been a number of studies documenting herb-layer 

response to fire, nearly all report short-term effects (<10 

years) of one to several fires, and 2) although the effects 

of fire vary among studies because of differences in 

vegetation and fire intensity, similar general responses 

were reported from many of the study sites.

THE HERBACEOUS LAYER

The herbaceous layer, also referred to as understory or 

groundlayer vegetation, generally is defined as all plants 

(woody and herbaceous) <1 m in height, though taller 

woody vegetation may be included (Gilliam and Roberts 

2003). Across oak forest landscapes, the herbaceous layer 

harbors the great majority of plant diversity, including 

1

USDA Forest Service, Northern Research Station, 359 

Main Road, Delaware, OH 43015, 740-368-0090, email: 

thutchinson@fs.fed.us.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 137

shallow soils) support woodlands as well as barrens 

communities usually containing unique species 

assemblages (Anderson et al. 1999). The intensity and 

frequency of past disturbances is a major factor related 

to current vegetation. For example, among secondary 

forests, those regenerated after agricultural conversion 

(old-field succession) have less diverse native herbaceous 

layers than continually forested stands (Flinn and 

Velland 2005). In many areas of the Eastern United 

States, deer browsing also has a major impact on herblayer vegetation (Côté et al. 2004).

Evidence indicates that periodic fire has occurred in 

forests for millennia (Patterson, this volume; Guyette, 

this volume; Ruffner, this volume). It has become clear 

that fire control has played an important role in the 

changing species composition of trees in oak forests, 

facilitating the abundant establishment of fire-sensitive 

and/or shade-tolerant tree and shrub species. Although 

it seems likely that more than 75 years of fire control 

also has had a large effect on herbaceous communities, 

long-term data to evaluate this hypothesis are lacking. 

However, a better understanding of fire effects on 

herbaceous vegetation is available from recent studies in 

which prescribed fire has been reintroduced to longunburned sites.

DIRECT FIRE EFFECTS ON PLANTS 

AND SEEDS

Most wildfires and prescribed fires in oak forests occur 

during the dormant seasons in late-winter/early spring 

and autumn (Haines and Johnson 1975) when most 

perennial herbs are dormant. Although the belowground rhizomes of most herbs are shallow (<10 cm), 

little heat is transferred into the mineral soil during 

typical prescribed fires (Boerner, this volume). In Ohio 

oak forests, the peak soil temperature during fires, at 

1-cm depth, averaged only 18o

C (64oF) (Iverson and 

Hutchinson 2002). Flinn and Pringle (1983) found 

that rhizome death of several forest understory species 

occurred only as temperatures approached 60oC (140oF). 

The fact that few perennial herbs decrease in abundance 

after fire, reported from a number of studies, suggests 

that their rhizomes are largely unaffected. However, the 

direct effects of fire on the rhizomes of oak forest herbs 

have not been investigated.

If spring fires are conducted later in the season when 

plants are emerging, direct heat damage to foliage 

occurs. For example, Primack et al. (1994) found that 

a late-spring fire reduced leaf area in a population of 

Cyprepedium acaule (pink lady’s slipper orchid) by 50 

percent; however, for plants with 10- to 50-percent leafarea damage, reproductive output returned to normal 

levels by the second year after fire.

In contrast to the perennial herbs, fire topkills the aboveground portions of nearly all small woody stems (shrubs, 

tree seedlings, vines). However, topkill induces sprouting 

(often vigorous) by hormonal signals, from dormant 

buds located below ground or on the base of stems. For 

example, Matlack et al. (1993) showed that fire increased 

stem density and shoot-growth of the shrub Gaylussacia 

baccata (black huckleberry).

Similar to fire effects on belowground rhizomes, fires 

likely cause little direct damage to seeds in the mineral 

soil. However, seeds within the litter and humus layers, 

which receive far more heat than mineral soil (Boerner, 

this volume) can be damaged by fire. Fire killed nearly 

50 percent of red oak acorns located in the litter layer 

(Auchmoody and Smith 1993).

In the early 1990s, it was first reported that smoke 

could directly stimulate seed germination (Brown and 

van Staden 1997). Since then, it has been shown to be 

a widespread phenomena, at least in fire-prone habitats. 

Of 215 species tested in the Cape Region of South 

Africa, 101 have exhibited increased germination when 

treated with smoke (Brown and van Staden 1997). In 

western Australia, smoke increased the germination 

of 45 of 94 species (Dixon et al. 1995). A butenolide 

compound derived from cellulose combustion (common 

to all vegetation fires) was responsible for the stimulation 

of germination by smoke (Flematti et al. 2004). The 

effects of smoke on germination have not been tested on 

plants native to the Eastern United States.

INDIRECT FIRE EFFECTS ON PLANTS 

AND SEEDS

Herb-layer vegetation is affected indirectly when fire 

alters the forest floor and soil environments. When fires 

are of low to moderate intensity and conducted in the 
1	 Fire 38 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

dormant season, these indirect effects arguably have a 

much greater effect on herb-layer vegetation relative to 

direct fire effects.

Leaf litter plays an important role in herb-layer plant 

communities (Facelli and Pickett 1991). In addition to 

its role in soil moisture and nutrient dynamics, leaf litter 

also serves as a mechanical barrier to seed germination 

and establishment. In the short term, the consumption 

of leaf litter and humus during fires probably causes the 

greatest changes in herb-layer vegetation by reducing the 

mechanical barrier and increasing light levels to seeds in 

the humus and mineral soil. Seeds of many herbaceous 

species in the eastern deciduous forest region have higher 

germination rates in light vs. dark conditions (Baskin 

and Baskin 1988). Throughout the growing season 

following spring fires, soil temperatures are elevated as 

the blackened litter absorbs more radiation (Iverson and 

Hutchinson 2002). Elevated soil temperatures likely 

affect the germination, establishment, and growth of 

herb-layer vegetation. The consumption of leaf litter in 

fires also releases nutrients that are then incorporated 

into the mineral soil, altering soil chemistry and likely 

affecting plant productivity, particularly on nutrientpoor sites (Gilliam and Christensen 1986; Boerner, 

this volume). If fire increases the amount of nitrate 

in soils, this could stimulate the germination of some 

seedbanking species (Auchmoody 1979).

As fire alters forest structure by topkill of saplings 

and some trees, light levels increase to the forest floor. 

However, in typical low-intensity fires, these changes 

are moderate and thus may have relatively little effect 

on plant performance. By contrast, high-intensity fires 

can open the canopy to a greater extent, increasing 

the competitive abilities of shade-intolerant herbs and 

shrubs. For example, in southern Ohio, all fires stimulate 

the germination of seedbanking, shade-intolerant species 

such as Erechtites hieracifolia (fireweed) and Rubus spp. 

(brambles) (Hutchinson et al. 2005a). After low-intensity 

fires when the canopy remains closed, these species rarely 

grow to more than several centimeters above the forest 

floor. However, when a portion of the canopy is removed 

by high-intensity fire or a combination of fire and 

harvesting, these species can rapidly grow to >1 m height 

in a single growing season (personal observation).

Fire has been shown to increase the reproduction of 

herbaceous species in several studies. In southeastern 

longleaf pine savannas, fire increased flowering (Platt et 

al. 1988) and clonal growth (Brewer and Platt 1994). 

For the rare plant Liatris scariosa var. novae-angliae

(northern blazing star), which occurs in grasslands in 

New England, Vickery (2002) found that prescribed fire 

reduced seed predation rates, from 90 percent before 

fire to 16 percent after fire. Little is known about the 

effects of fire on the flowering and seed production of 

herbaceous species in oaks forests.

FIRE EFFECTS ON HERB-LAYER 

COMMUNITIES: THREE CASE

STUDIES

It is difficult to determine the effects of fire on herb 

layer communities because of differences in vegetation 

and fires among studies. Therefore, to illustrate specific 

fire effects, I will first summarize the results of three 

studies. Each study: 1) had a duration of at least 5 years, 

2) involved repeated dormant-season fires that reduced 

understory and midstory tree density but had little affect 

on overstory trees, and 3) included both pretreatment 

and multiple years of postreatment plant community 

data.

Missouri Ozarks Dry Oak Forest 

(Hartmann and Heumann 2003)

The study was conducted in the dissected 1,012-ha 

Chilton Creek basin, owned and managed The Nature 

Conservancy. The presettlement landscape consisted of 

open woodlands dominated by shortleaf pine and white 

oak. Early descriptions suggest an understory that was 

dominated by grasses and forbs. Postsettlement logging, 

grazing, and fire suppression have greatly altered the 

landscape, which now consists of closed-canopy oak 

forests averaging 13 m2

/ha of basal area. Although the 

area contains more than 500 herbaceous species, woody 

plants and hardwood leaf litter dominate the herb-layer 

cover. The authors state that fire is being applied to 

create a “mosaic of high quality native habitats.” Five 

separate units were burned 1 to 4 times (1998-2001). 

Herb-layer data were collected in 250 plots (4,000 1-m2

quadrats) prior to fires, and again in 1998 and 2001. In 

all, 486 species (99 percent native) were recorded.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 139

By 2001, forest structure had become somewhat 

more open on the burned units (all five combined) as 

understory (1 to 4 cm d.b.h.) and midstory (4 to 11

cm d.b.h.) stems were reduced by 47 and 28 percent, 

respectively. Fires increased the cover and frequency 

of herbaceous plants relative to woody plants. Among 

herbs, the cover of legumes more than doubled to 8 

percent, and the cover of other forbs, grasses, and sedges 

all increased over pretreatment levels, but each remained 

at less than 5 percent total cover. Comon herbaceous 

species that increased in relative importance included 

Panicum boscii (Bosc’s panic grass), Carex nigromarginata

(black-margined sedge), Brachelytrum erectum (longawned wood grass), Helianthus hirsutus (woodland 

sunflower), and Lespedeza intermedia (wand lespedeza). 

Total vegetation cover in the herb layer increased from 

15 to 24 percent on burned sites while hardwood litter 

cover was reduced by 30 percent. Over 5 years, smallscale species richness increased slightly on burned sites 

(11.5 to 12.2 species/1 m2

) as did the total number of 

species recorded each year (465 to 482).

The authors concluded that the prescribed fires had 

created a “landscape in transition” as the understory 

was more open but the herb-layer response had been 

moderate. They hypothesized that the longer-term 

application of periodic fire (20+ years) seems necessary to 

restore the area to a “more open and biologically diverse 

landscape.”

Ohio Dry-Mesic Oak Forest

(Hutchinson et al. 2005a)

Our study was conducted on four 75-ha sites in 

the dissected Allegheny Plateau of southern Ohio. 

Presettlement forests were dominated by oak. Currently, 

second-growth forests that established prior to fire 

suppression (ca. 1850 to 1900) remain oak-dominated, 

but shade-tolerant trees (e.g., red maple) are abundant 

in the midstory and understory. Tree basal area averaged 

28 m2

/ha. Fire treatments from 1996-99 were no fire, 

burned 2X, and burned 4X. Species’ frequencies were 

recorded annually in 108 plots (1,728 2-m2

 quadrats) 

over a 5-year period (1995-99). Plots were stratified by 

an integrated moisture index into xeric, intermediate, 

and mesic classes. Overall, 452 species (97 percent 

native) were recorded.

Fires reduced the density of saplings (<10 cm d.b.h.) by 

more than 80 percent. The 2X and 4X burn treatments 

produced similar results. Species composition was 

significantly affected by fire but differences between 

unburned and burned sites were relatively minor 

compared to compositional differences between dry and 

mesic sites. Species composition shifted on all moisture 

classes after fire, but to a greater degree on dry plots. 

Total species richness on burned sites increased from 

17.1/2 m2

 before fires (1995) to 18.5/2 m2, averaged 

across post-burn years. On unburned plots, richness 

decreased slightly from 15.3 to 15.1/2 m2

. 

Among species groups, richness increased significantly 

for annual forbs, summer-season perennial forbs, grasses, 

and woody seed-banking species, while tree-seedling 

richness decreased. Perhaps the most striking result was 

that the vast majority of perennial forb species exhibited 

little change in frequency on burned sites over the 5-year 

study (Appendix A). Only 7 of 49 common forb species 

exhibited a change in frequency of ≥ 5 percent; only one 

of those, Viola spp. (Violets), changed more than 10 

percent, increasing on burned units from a mean of 30.5 

percent before fire to 49.2 percent on the burn sites after 

5 years.

In a separate posttreatment sample of 480 1-m2

 quadrats 

located in six 2-ha stands within the same study sites (3 

years after the last fire), herbaceous cover averaged 17.1

percent in burned sites vs. 4.7 percent in unburned sites 

(Hutchinson 2004). For each of the major herbaceous 

species groups (grasses, sedges, legumes, composits), 

cover was higher on the burned sites (Fig. 1); the 

combined cover of the these four groups averaged 

12.3 and 3.3 percent in burned and unburned stands, 

respectively.

Southern Illinois Oak Barrens 

(Taft 2003)

This study was conducted on a 3-ha dry sandstone 

barrens owned by The Nature Conservancy, and also 

on a nearby unburned barrens in the Shawnee National 

Forest. Both barrens were dominated by post oak 

(Quercus stellata). Tree basal area averaged 18 m2/ha. 

Herb-layer cover was high at both sites (>90 percent) 

prior to treatment. Fire was applied to the barrens 
1	 Fire 40 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

to prevent its conversion to dry forest and sustain 

and enhance the herbaceous flora of this rare natural 

community. Fires were conducted in 1989 (fall) and 

1994 (spring). Herb-layer vegetation was sampled in 

1989 (pretreatment) and four posttreatment years. 

Twenty-three plots (276 0.25-m2

 quadrats) were sampled 

each year.

Density of stems in the shrub/sapling layer (50 cm 

height to < 6 cm d.b.h.) was reduced by 55 percent on 

the burned site after fire. Herbaceous cover increased 

from 90 to 117 percent on the burned sites and 

concurrently decreased from 98 to 61 percent on the 

unburned site. Small-scale species richness more than 

doubled on the burned site, to > 8 species/0.25 m2

 while 

remaining static on the unburned site. The total number 

of species sampled increased from 94 to 121 on the 

burned site and decreased from 74 to 68 species on the 

unburned site.

Most of the species that increased on the burned site 

were perennial forbs, grasses (e.g., several Panicum 

species, the most abundant of which was P. laxiflorum,

pale green panic grass), and sedges typical of dry 

woodland habitats. The most common mode of 

establishment after fire was germination of seed 

stored in the soil. The only two herbaceous species to 

decrease after fire were, surprisingly, the prairie grasses 

Schizachyrium scoparium (little bluestem) and Sorgastrum 

nutans (indian grass), presumably because light remained 

limiting to their productivity.

The author concluded that periodic fire (every 3 to 4 

years) is necessary to maintain the composition and 

diversity of the barrens flora. Without fire to stimulate 

the seedbank, some less common species likely would 

become locally extinct as woody succession proceeds.

FIRE EFFECTS ON HERB 

COMMUNITIES: GENERALIZATIONS

When observing a typical oak forest that has been 

burned recently, the most noticeable effects of fire are: 

1) a more open structure caused by topkill of the sapling 

and shrub layer, 2) the prolific sprouting of most woody 

plants, and 3) the greater cover of herbaceous plants, 

including forbs, grasses, and sedges. Nearly all studies of 

fire effects, in addition to those described earlier, have 

revealed that the cover and/or abundance of herbaceous 

plants increased after fire (e.g., Swan 1970; McGee et al. 

1995; Nuzzo et al. 1996; Arthur et al. 1998). For woody 

plant cover, the fire response may be more variable 

because of the variation in the sprouting response in 

different vegetation types (e.g., Nuzzo et al. 1996; Elliot 

et al. 1999; Kuddes-Fischer and Arthur 2002). The 

abundance of woody stems often increases because of 

sprouting (e.g., Ducey et al. 1996; Arthur et al. 1998), 

and foliage cover is shifted from the midstory/understory 

to the herb layer.

Figure 1.—Photograph (2005) of a ridgetop 

site in southern Ohio that was burned 

annually 1996-1999 and again in 2004. Here 

the overstory is dominated by white oak, the 

sapling layer has largely been eliminated, 

and common species in the herbaceous layer 

include Panicum commutatum, Panicum 

boscii, and seedlings of sassafras and oak. 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 141

As with the three case studies, fire often has been shown 

to increase species richness and/or diversity in the 

herbaceous layer (e.g., Arthur et al. 1998; Nuzzo et al. 

1996; Elliot et al. 1999), though results vary among 

studies. By contrast, Ducey et al. (1996) showed that 

diversity was reduced after fire in some areas, likely due 

to abundant sprouting from Kalmia latifolia (mountain 

laurel). Several studies have shown no significant effects 

of fire on herb-layer diversity (Luken and Shea 2000; 

Kuddes-Fischer and Arthur 2002; Franklin et al. 2003).

The increases in small-scale species richness after fire 

largely result from species already present on the site 

that increase via germination after fire. Species groups 

that commonly respond to fire include woodland grasses 

(e.g., Panicum spp.), sedges, composites, legumes (e.g., 

Desmodium spp, Lespedeza spp.), and woody seedbanking 

species (e.g., Liriodendron tulipifera, Rubus spp.).

In southern Ohio we have documented the 

establishment of several species on burned sites that were 

absent or rare before fire, including Ceanothus americanus

(New Jersey Tea), Phaseolus polystachios (wild kidney 

bean), Clitoria mariana (butterfly pea), Rhus glabra

(smooth sumac), Chamaecrista nictitans (wild sensitiveplant) Rhus copallinum (winged sumac), Desmodium 

cuspidatum (large bract tick trefoil), Eupatorium 

sessilifolium (upland boneset), Eupatorium serotinum

(late-flowering thoroughwort), Phytolacca americana

(pokeweed), Hackelia virginiana (common stickseed), 

Lobelia inflata (indian-tobacco), Sphenopholis nitida

(shining wedge grass), and Helianthus microcephalus

(small wood sunflower). Although there is the potential 

for invasive exotic plants to establish or to increase 

in abundance after fire (Huebner, this volume), most 

studies have reported only minor changes in the 

abundance exotic plants.

FIRE EFFECTS ON HERB 

COMMUNITIES: CAUSES OF 

VARIATION

Fire intensity can vary dramatically both among different 

fires and across the landscape within a fire. Although 

most prescribed fires have relatively little affect on 

overstory trees, intense surface fires can cause significant 

overstory mortality (Regelbrugge et al. 1994; Ducey et 

al. 1996; Elliot et al. 1999). In Connecticut oak forests 

with abundant Kalmia latifolia (mountain laurel), Ducey 

et al. (1996) found that postburn plant diversity was 

higher in intensely burned patches, where the overstory 

had been killed, than moderately burned patches, where 

Kalmia was the most dominant after sprouting.

In landscapes with significant topographic heterogeneity, 

vegetation also varies across the landscape. In the 

southern Appalachians, Elliot et al. (1999) found that 

prescribed fire intensity and effects on the overstory 

and understory increased along an upslope moisture 

gradient from a mesic cove community to dry midslope 

oak community to a xeric ridge pine-oak community. 

In southern Ohio, where total elevation changes are 

less pronounced than in the southern Appalachians, we 

found that fire intensity and effects were less pronounced 

from dry to mesic sites (Hutchinson et al. 2005a). 

In other ecosystems, fire has reduceed differences in 

vegetation across topographic gradients (e.g., Gibson and 

Hulbert 1987; tallgrass prairie) or reinforced differences 

(Liu et al. 1997, longleaf pine forest).

Fire-season effects on herb-layer vegetation seldom have 

been compared in oak forests. In a dry-mesic “degraded” 

forest in northern Illinois, Schwartz and Heim (1996) 

found that the abundance and richness of native herbs 

was reduced for several years by a May growing-season 

fire but not by a March dormant-season fire. In restored 

shortleaf pine grassland communities in Arkansas, 

Sparks et al. (1998) compared late dormant-season burns 

(March-April) to late growing-season burns (SeptemberOctober) and found that differences were relatively 

minor. However, March and July burns produced 

different vegetation responses for some early-season and 

late-season species in planted tallgrass prairies (Howe 

1995). In longleaf pine savannas, growing-season burns 

are more effective at suppressing midstory deciduous 

trees (Quercus spp.) and understory shrubs than 

dormant-season fires (Glitzenstein et al. 1995; Drewa et 

al. 2002).

Not surprisingly, different fire frequencies applied over 

a short period have produced largely similar herb layer 

vegetation responses in oak forests (e.g., Hutchinson 

et al. 2005a). However, in other vegetation types 
1	 Fire 42 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

where different fire frequencies have been applied over 

several decades, vegetation response can be significantly 

different. For example, long-term differences in 

fire frequency at Konza Prairie in Kansas indicate 

that frequent burning (annual or biennial) tends to 

homogenize vegetation by favoring the dominant warmseason grass Andropogon gerardii (big bluestem) to a 

much greater extent than infrequent burning (Collins 

1992).

When prescribed fire has been applied to more openstructured communities within oak forest landscapes in 

which woodland and/or prairie species have persisted, 

fire effects usually are more pronounced compared to fire 

effects in closed-canopy oak forests. In addition to the 

oak barrens study described (Taft 2003), Nuzzo (1996) 

reported a 50-percent increase in herb-layer richness after 

prescribed fires in a low-density Illinois “sand forest.” In 

a 25-year study of fire effects in a Tennessee oak barrens, 

unburned plots showed a sharp decline in herbaceous 

cover as woody succession proceeded, while burned plots 

maintained high coverage of both forbs and graminoids 

throughout (DeSelm and Clebsch 1991).

EFFECTS OF SILVICULTURE AND FIRE

ON THE HERBACEOUS LAYER

Although prescribed fire is being applied more often to 

sustain oak forests, research has shown that fire alone, 

at least in the short term, often does not improve the 

competitive status of oak regeneration because the 

canopy remains closed and competing species also sprout 

readily (Brose, this volume; Hutchinson et al. 2005b). 

The combined use of silvicultural treatments that remove 

a portion of the canopy followed by prescribed fire has 

improveed oak regeneration in some cases (Kruger and 

Reich 1997; Brose and Van Lear 1998).

The effects of timber harvesting on herb communities 

has received much attention, some of which has been 

controversial. Although results differ substantially among 

studies (Roberts and Gilliam 2003), some of the more 

thorough studies have shown no reduction in herb layer 

diversity following timber harvest, either in the short 

term (e.g., Grabner and Zenner 2002) or in the longer 

term (e.g., Elliot et al. 1997).

Few studies have examined the effects of fire, harvesting, 

and combinations of the two on herb-layer vegetation in 

oak forests. The national Fire and Fire Surrogates Study 

is quantifying the effects of fire and other treatments 

that alter forest structure (e.g., mechanical thinning) at 

13 sites across the country. At the Ohio Hills Site, we 

are monitoring the response of herb layer communities 

to fire only, thinning only, and thinning followed by 

fire. Preliminary data indicate clear differences in herblayer response to thinning and fire treatments (Fig. 2). 

Species richness and herbaceous cover have increased 

much more after fire (and thinning+fire) than after 

Figure 2.—Response of herbaceous layer vegetation to 

thinning, burning, and thinning+burning treatments at two 

replicate sites (Raccoon Ecological Management Area 

and Zaleski State Forest) of the Ohio Hills Fire and Fire 

Surrogate study site. Mean annual values (± 1 S.E.) for a) 

species richness per 1 m2 quadrat, b) the combined foliar 

cover (percent) of all herbaceous species <1 m above the 

forest floor, and c) combined foliar cover (percent) of all 

woody species <1 m above the forest floor. In each graph, 

the dashed line shows the timing of both thinning and 

burning treatments: thinning occurred from autumn 2000 to 

spring 2001; fires were conducted in March and April 2001. 

Herb-layer data were collected in the summers (JuneAugust) of 2000, 2001, and 2004.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 143

thinning alone. By contrast, the total cover of woody 

plants (shrubs and tree seedlings+sprouts), has increased 

substantially on all treatments, relative to untreated 

stands. These results suggest that fire as a process is a 

unique disturbance that may be required to stimulate 

the seedbank and increase the diversity and productivity 

of the oak-forest herbaceous layer. Two other short-term 

studies of thinning and fire have shown less substantial 

responses of herb-layer vegetation (Franklin et al. 2003; 

Dolan and Parker 2004).

RARE PLANTS AND FIRE

Owen and Brown (2005) divided the 186 federally listed 

plant species into four categories of fire adaptation and 

tolerance: 47 species require fire, 65 tolerate fire, 70 

occur in habitats where fire does not occur (e.g., aquatic 

plants), and are adversely affected by fire.

Throughout the central hardwoods region, prescribed 

fire is used to sustain and restore open-structured plant 

communities (e.g., barrens, woodlands) that usually 

support rare species. However, maintaining rare plant 

populations is seldom the primary objective of fire use 

in oak forests. Some rare plants that occur in upland oak 

forests, particularly those threatened by shading, could 

potentially benefit from prescribed fire through increased 

germination, establishment, and reproduction. In Ohio, 

I compiled a list of 125 state-listed rare plant species 

that occur primarily in the “hill country” of southern 

Ohio. I excluded species that occur in habitats where fire 

does not occur. For each species, the Ohio Department 

of Natural Resources, Division of Natural Areas lists 

13 primary threats to the populations (http://www.dnr.

state.oh.us/dnap/heritage/Rare_Species2004.html). By 

far the most common threat among the 125 species is 

“shading as a result of woody plant succession,” which 

applies to 71 species. Prescribed fire is a potential tool for 

maintaining the open habitats required by these species. 

However, for some of these species, other methods (e.g., 

mechanical removal of trees, herbicide application, 

mowing) could be more effective or achieved more 

readily than burning.

For nearly all rare plants that occur in oak forest 

landscapes, the effects of fire are unknown. In southern 

Ohio, I quantified the response of the state-endangered 

Calamagrostis porterii subsp. insperata (Bartley’s bent reed 

grass) that occurs as distinct clonal patches, primarily on 

dry-mesic ridgetops in oak forests (Hutchinson 2004). 

Tiller density increased in patches burned annually from 

1996 to 1999 compared with unburned patches, though 

cover and patch area were unaffected. Annual fires also 

stimulated flowering, which has been observed only 

infrequently in natural settings. From 1995 to 2001, 125 

flowering stems were documented on the seven annual 

burn patches (total area of patches = 229 m2

) compared 

to only 13 flowering stems on the 33 unburned patches 

(area = 999 m2).

MANAGEMENT IMPLICATIONS

The use of prescribed fire to restore oak-forest structure, 

and to improve oak regeneration and wildlife habitat is 

a critically important management issue, as evidenced 

by the nearly 400 attendees at this conference. As the 

use of prescribed fire in oak forests is expanding rapidly, 

it is important to better understand its effects on other 

ecosystem components such as the diverse herbaceous 

layer. Research indicates that typical prescribed fires, 

those conducted during the dormant season and of 

low to moderate intensity, can be applied to oak forests 

without having a major impact on the herbaceous 

layer vegetation. The increase in herbaceous diversity 

and cover, documented after fire in most studies, is 

desirable to many land managers. While the implications 

of fire management are perhaps less obvious to the 

herbaceous layer of oak forests than in more open 

communities, such as barrens and woodlands, the fireinduced germination of some species and the more open 

conditions promoting their growth and reproduction, 

likely are important for the long-term maintenance of 

species diversity. Local populations of some rarer species 

may be threatened by the continued exclusion of fire.

Although research has shown general patterns of 

herb-layer response to fire, the implications of fire 

management likely will vary substantially across 

the central hardwoods region and also within local 

landscapes. To promote landscape-scale plant diversity, it 

may be desirable to apply periodic fire to dry upland sites 

while continuing to exclude fire from mesic sites. This 

approach is being used in the southern Appalachians 
1	 Fire 44 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

where ridgetop ignitions of large units result in fires that 

burn completely and at moderate to high intensity in 

xeric uplands, contrasting with a patchy mosaic of lowintensity and unburned areas in mesic low slopes (Hugh 

Irwin, personal communication).

Studies of fire in oak forests have not shown significant 

establishment or increased abundance of invasive exotic 

plant species (Huebner, this volume). However, because 

many invasives are adapted to disturbed conditions 

for germination and growth, fire-induced alterations 

to the forest floor and canopy could facilitate their 

establishment, particularly if invasives are common 

and fires are of high intensity and/or combined with 

harvesting. Prior to burning, invasives should be treated 

to reduce the likelihood of postburn establishment. The 

abundant establishment of invasive species after fire 

almost surely would have a negative impact on native 

herbaceous communities.

ACKNOWLEDGMENTS

I thank Stephen Brewer and Tim Nigh for reviewing 

a previous draft of this manuscript, Pat Brose and the 

organizers of the conference for the invitation to make 

the presentation, Matthew Dickinson for editorial 

advice, and John Taft and Blane Heuman for providing 

information and photographs of their study sites in 

Illinois and Missouri, respectively. The Ohio prescribed 

fire study was supported by a grant to Elaine Kennedy 

Sutherland from the USDA Forest Service’s program 

on Ecosystem Management Research. The Fire and Fire 

Surrogate Study in Ohio was supported by a grant from 

the Joint Fire Science Program.

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ecology: six pragmatic models for understanding 

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96: 177-189.

Kruger, E. L.; Reich, P. B. 1997. Responses of 

hardwood regeneation to fire in mesic forest 

openings. I. Post-fire community dynamics.

Canadian Journal of Forest Research. 27: 1822-1831.

Kuddes-Fischer, L.M.; Arthur, M.A. 2002. Response of 

understory vegetation and tree regeneration to a 

single prescribed fire in oak-pine forests. Natural 

Areas Journal. 22: 43-52.

Liu, C.; Harcombe, P.A.; Knox, R.G. 1997. Effects of 

prescribed fire on the composition of woody plant 

communities in southeastern Texas. Journal of 

Vegetation Science. 8: 495-504.

Luken, J.O.; Shea, M. 2000. Repeated prescribed 

burning at Dinsmore Woods State Nature Preserve 

(Kentucky, USA): responses of the understory 

community. Natural Areas Journal. 20: 150-158.

Matlack, G.R.; Gibson, D.J.; Good, R.E. 1993. 

Regeneration of the shrub Gaylussacia baccata

and associated species after low-intensity fire in an 

Atlantic coastal plain forest. American Journal of 

Botany. 80: 119-126.

McGee, G.G.; Leopold, D.J.; Nyland, R.D. 1995. 

Understory response to springtime prescribed 

fire in two New York transition oak forests. Forest 

Ecology and Management. 1: 149-168.

Nuzzo, V.A.; McClain, W.; Strole, T. 1996. Fire impact 

on groundlayer flora in a sand forest 1990-1994.

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Owen, W.; Brown, H. 2005. The effects of fire on rare 

plants. Fire Management Today. 65: 13-15.

Platt, W.J.; Evans, G.W.; Davis, M.M. 1988. Effects 

of fire season on flowering of forbs and shrubs in 

longleaf pine forests. Oecologia. 76: 353-363.
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Primack, R.B.; Miao, S.L.; Becker, K.L. 1994. Costs 

of reproduction in the pink lady’s slipper orchid 

(Cypripedium acaule): defoliation, increased fruit 

production, and fire. American Journal of Botany. 

81: 1083-1090.

Regelbrugge, J.C.; Smith, D.W. 1994. Postfire tree 

mortality in relation to wildfire severity in mixed 

oak forests in the Blue Ridge of Virginia. Northern 

Journal of Applied Forestry. 11: 90-97.

Schwartz, M.W.; Heim, J.R. 1996. Effects of a 

prescribed fire on degraded forest vegetation.

Natural Areas Journal. 16: 184-191.

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Bukenhofer, G.A. 1998. Effects of late growingseason and late dormant-season prescribed fire on 

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communities. Journal of Vegetation Science. 9: 133-

142.

Swan, F.R. 1970. Post-fire response of four plant 

communities in south-central New York state.

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barrens. Journal of the Torrey Botanical Society. 130. 

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rare grassland perennial. Conservation Biology. 16; 

413-421.
1	 Fire 48 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Appendix

Mean frequency of common species over 5 years on burn units on four study sites in southern Ohio. Each 

site contained a unit burned 2X (1996 and 1999) and a unit burned 4X (1996 to 1999); here the burn 

treatments have been combined. Each year sixteen 2-m2

 quadrats were sampled within each of nine 625-m2

plots per burn unit (total of 72 plots).

Frequency (%)

1995 1996 1997 1998 1999

Scientific name Common name (preburn)

Graminoids

 Brachyelytrum erectum long-awned wood grass 21.3 22.2 25.3 22.4 27.2

 Bromus pubescens woodland brome 6.5 3.8 4.8 4.8 5.4

 Carex digitalis narrow-leaved wood sedge 9.9 6.1 6.3 10.0 6.7

 Carex gracilescens slender wood sedge 10.9 9.6 13.6 12.0 15.0

 Carex rosea stellate sedge 4.5 4.0 3.4 4.2 4.6

 Carex spp. sedge spp. 6.9 27.4 17.6 22.1 37.6

 Carex wildenovii Willdenow’s sedge 12.8 5.7 10.2 14.0 10.9

 Danthonia spicata poverty oat grass 4.6 2.3 3.6 5.4 5.7

 Diarrhena americana beak grass 2.7 3.3 2.8 3.0 3.8

 Panicum boscii Bosc’s panic grass 15.1 14.9 21.6 25.0 29.8

 Panicum commutatum variable panic grass 0.7 7.0 20.7 22.0 23.1

 Panicum dichotomum forked panic grass 6.2 7.9 10.2 12.5 13.3

 Poa cuspidata cuspidate spear grass 16.8 13.3 16.4 12.8 12.5

Annual forbs

 Acalypha virginica three-seeded mercury 1.5 10.9 10.9 11.7 18.2

 Amphicarpaea bracteata hog-peanut 20.9 23.1 26.2 25.9 21.5

 Erechtites hieracifolia fireweed 5.9 72.7 35.9 30.5 34.8

 Galium aparine cleavers 12.3 3.1 10.6 8.3 0.7

 Pilea pumila common clearweed 7.9 10.1 9.8 11.2 10.9

Perennial forbs

 Agrimonia spp. agrimony spp. 5.9 5.1 7.6 6.5 5.7

 Anemonella thalictroides rue anenome 24.7 24.3 25.7 25.7 25.6

 Arisaema triphyllum jack-in-the-pulpit 19.1 17.7 17.4 19.0 18.6

 Aristolochia serpentaria Virginia-snakeroot 10.3 12.5 12.2 11.3 11.9

 Asarum canadense wild ginger 13.5 10.5 11.5 11.9 10.1

 Aster divaricatus white wood aster 10.3 9.7 12.3 13.9 10.1

 Aureolaria laevigata entire-leaved false foxglove 2.2 2.3 2.5 3.7 4.7

 Cardamine angustata slender toothwort 6.8 3.0 7.0 6.2 6.3

 Chimaphila maculata spotted pipsissewa 4.9 4.2 4.1 4.6 0.7

 Cimicifuga racemosa black cohash 14.7 15.7 16.0 15.0 16.2

 Circaea lutetiana c. enchanter’s nightshade 11.1 8.9 9.9 10.7 10.6

 Collinsonia canadensis richweed 3.2 5.6 5.2 5.6 5.4

 Conopholis americana squaw-root 3.6 3.9 2.9 4.4 4.5

 Cunila oreganoides common dittany 4.1 3.6 4.2 4.4 5.1

 Desmodium glutinosum cluster-leaved tick-trefoil 11.5 12.2 10.6 12.4 11.4

 Desmodium nudiflorum naked tick-trefoil 43.8 43.1 42.4 46.3 38.7

 Dioscorea quaternata wild yam 12.5 11.9 12.0 13.3 12.2

 Eupatorium rugosum white snakeroot 14.8 19.8 23.9 24.9 22.7

 Galium circazans wild licorice 27.8 21.3 28.6 31.3 31.9

 Galium concinnum shining bedstraw 6.4 6.5 7.2 6.9 7.6

 Galium triflorum sweet-scented bedstaw 30.3 42.4 43.8 44.0 37.4

 Geranium maculatum wild geranium 35.8 33.9 31.1 31.3 29.8

 Geum spp. avens spp. 3.1 3.1 4.6 3.4 3.3
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 149

 Helianthus divaricatus woodland sunfower 5.5 10.8 8.0 7.4 6.8

 Hieracium venosum veined hawkweed 4.3 3.1 3.2 4.3 3.7

 Hydrophyllum macrophyllum large-leaved waterleaf 5.3 4.2 4.6 4.6 5.0

 Lespedeza spp. lespedeza spp. 1.5 4.8 11.6 7.9 11.2

 Lysimachia quadriflora whorled loosestrife 7.0 7.8 8.6 12.1 10.3

 Medeola virginiana indian cucumber-root 3.2 2.8 3.6 3.9 3.3

 Monarda fistulosa wild bergamot 5.1 4.2 5.6 5.8 4.4

 Osmorhiza claytonii wooly sweet cicely 9.1 6.2 6.1 6.2 4.4

 Oxalis violacea violet wood-sorrel 2.6 3.9 2.9 4.6 4.5

 Phlox divaricata blue phlox 6.2 7.3 6.3 4.9 4.2

 Podophyllum peltatum mayapple 9.7 8.7 8.9 8.7 8.1

 Polygonatum biflorum smooth solomon’s seal 13.6 6.0 13.5 9.9 12.9

 Polygonum virginianum jumpseed 6.5 10.2 8.2 7.1 8.6

 Potentilla spp. cinquefoil 14.4 13.7 18.8 19.5 19.5

 Prenanthes spp. rattlesnake root 14.6 15.5 15.5 14.3 11.5

 Sanguinaria canadensis bloodroot 6.6 5.8 7.3 8.2 6.9

 Sanicula spp. snakeroot 16.1 12.4 18.8 16.9 15.9

 Scutellaria spp. skullcap 10.2 10.8 10.5 11.7 11.8

 Smilacinia racemosa false solomon’s seal 31.4 15.6 29.7 26.7 28.0

 Solidago caesia blue-stemmed goldenrod 13.5 16.8 14.1 17.4 16.3

 Solidago flexicaulis zigzap goldenrod 3.2 5.4 4.9 5.0 3.7

 Tiarella cordifolia foamflower 11.3 10.5 11.6 10.9 10.2

 Trillium grandiflorum large white trillium 16.5 7.5 16.8 15.1 16.7

 Uvularia perfoliata peroliate bellwort 38.3 28.9 34.5 35.8 36.1

 Vicia caroliniana pale vetch 3.0 5.0 5.1 4.3 2.6

 Viola spp. violets 30.5 52.2 55.2 55.1 49.2

Pteridophytes

 Adiantum pedatum maidenhair fern 3.3 3.3 3.9 3.3 3.3

 Botrychium virginianum rattlesnake fern 10.3 4.9 10.5 8.4 9.2

 Osmunda claytonia interrupted fern 1.5 1.6 2.3 2.0 1.9

 Polystichum acrosticoides Christmas fern 19.5 19.0 21.1 20.4 21.4

 Thelypteris hexagonoptera broad beech fern 3.6 4.3 4.5 4.8 4.5

Shrubs

 Corylus americana American hazel 4.6 5.8 4.2 4.7 3.0

 Hamamelis virginiana witch-hazel 4.8 4.3 4.8 3.8 3.6

 Hydrangea arborescens wild hydrangea 7.5 7.1 8.0 8.9 7.6

 Lindera benzoin spicebush 17.3 15.7 16.5 16.2 12.8

 Rhus glabra smooth-sumac 0.0 12.0 4.1 6.4 13.9

 Rosa carolina pasture rose 14.1 13.9 12.5 13.9 13.1

 Rubus spp. brambles 22.1 28.3 41.1 40.2 43.5

 Smilax glauca sawbrier 27.6 28.0 27.3 29.1 30.9

 Smilax hispida bristly greenbriar 5.7 6.7 5.2 6.4 5.6

 Smilax rotundifolia common greenbriar 41.0 39.4 37.2 36.5 34.5

 Vaccinium palidum low blueberry 21.0 22.0 21.8 21.3 21.3

 Vaccinium stamineum deerberry 7.1 3.0 4.0 3.1 2.7

 Viburnum acerifolium maple-leaved viburnum 23.3 19.8 19.6 19.5 17.8

Vines

 Parthenocissus quinquifolius Virginia creeper 38.5 36.9 32.9 33.1 25.7

 Toxicodendron radicans poison ivy 15.7 22.2 9.5 10.9 8.2

 Vitis sp. wild grapvine 21.3 60.3 47.8 42.4 37.4

Appendix—continued
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Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 151

Fire - Fauna Interactions
1	 Fire 52 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

DO FIRE AND INSECTS INTERACT IN EASTERN FORESTS?

Lynne K. Rieske-Kinney1

Abstract.—The increasing use of prescribed fire as a management strategy for manipulating forest-species 

composition generates questions regarding the effects on the arthropod community and the underlying 

processes in which arthropods play a dominant role, as well as its potential as a pest suppression strategy. 

Despite the apparent benefits of prescribed burning for manipulating stand composition and enhancing 

tree vigor, relatively little is known about how fire interacts with arthropod-dependent processes in 

eastern forest ecosystems. This paper reviews the evidence of direct and indirect interactions between 

forest arthropods and fire, and addresses the following questions: 1) are soil- and litter-dwelling 

arthropods irreparably harmed by burning? 2) does prescription burning alter plant susceptibility to 

insect herbivores? 3) can fire be used as a management strategy to suppress forest arthropod pests?

Although soil- and litter-dwelling arthropod abundance is affected by prescribed burning, arthropod 

diversity and richness are not. Litter arthropod evenness increases in response to burning, most 

likely due to reductions in mites and collembolans, the two dominant taxa. Fire-induced changes 

in foliar chemistry often are transient and may be species-specific. These changes are not fully 

predictable but could alter patterns of insect herbivory. Use of prescribed fire for pest suppression 

in managed forests has lagged behind that of other managed systems, and it has had limited use 

for pest suppression in deciduous forests of the Eastern United States. The highly clustered spatial 

distribution of acorn predators makes effective suppression through prescription burning problematic.

Insects and related arthropods are essential to forest 

ecosystem processes. Arthropods dominate forest 

soils and leaf litter, and play vital roles in litter 

decomposition, nutrient dynamics, soil development, 

and soil stability (Wood 1995). Arthropods contribute to 

decomposition of organic matter and soil development 

by reducing the size of organic particles, consequently 

accelerating fungal and bacterial decomposition (Metz 

and Dindal 1975). Arthropod fungivores also stimulate 

fungal growth by hyphal grazing, thereby influencing 

the balance between fungi and bacteria (Hanlon and 

Anderson 1979).

As herbivores, arthropods also play critical roles in 

ecosystem processes. Herbivores can function as 

seed predators, causing extensive, localized loss of 

regeneration in eastern oak forests (Drooz 1985). 

Acorn predators such as acorn weevils, sap beetles, and 

the acorn moth are primary and/or secondary pests 

(feeding and breeding in intact versus damaged seed), 

consuming acorn cotyledons and damaging radicles, or 

directly damaging germinated seedlings (Van Leeuwen 

1952; Gibson 1964; Galford and Weiss-Cottrill 1991; 

Galford et al. 1988, 1995). Acorn predators are closely 

associated with soil and litter and as such may vector 

INTRODUCTION

Prescribed fire is increasingly used as an intermittent 

disturbance agent to mimic pre-settlement disturbance 

regimes. This management strategy is designed to 

encourage regeneration and stand development (Lorimer 

1993). Prescribed fire combusts the leaf litter and can 

reduce total microbial and fungal biomass (Fritze et 

al. 1994). Burning can lead to increased soil pH and 

greater fluctuations in temperature and moisture, with 

subsequent loss of vegetation (Haimi et al. 2000). 

Burning suppresses vegetative competition and enhances 

light penetration to the forest floor. In oak-dominated 

forests of the Eastern United States, fire helps reduce 

invasive fire-sensitive maples (Arthur et al. 1998), and 

makes conditions more favorable for development and 

growth of oak seedlings (Reich et al. 1990; Adams and 

Rieske 2001). The increasing use of prescribed fire as 

a management strategy for manipulating forest-species 

composition generates questions regarding the effects 

on the arthropod community and on the underlying 

processes in which arthropods play a vital role.

1

University of Kentucky, Department of Entomology, 

Lexington, KY 40546-0091, 859-257-1167, email: lrieske@

uky.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 153

and/or create infection courts for potentially pathogenic 

microorganisms. Additionally, these arthropods are 

strongly affected by factors such as stand composition, 

litter quality and depth, and soil type and depth, which 

influence their local distribution and impact (Dindal 

1990).

Defoliating herbivores alter net primary productivity, 

increasing nitrogen input and light penetration. 

Indirectly, defoliators can influence watershed 

characteristics and, through changes in resource 

availability, affect wildlife distribution patterns. 

Herbivore population outbreaks cause widespread 

mortality that can shift forest-stand composition and 

influence ecological succession (Drooz 1985). Insect 

herbivory is one of a suite of disturbance factors 

that have contributed to the development of today’s 

forests. In addition to herbivore pressure, intermittent 

disturbances consisting of pathogens, anthropogenic 

activities, climactic pressures, and fire have influenced 

the formation and maintenance of forests of Eastern 

North America (Abrams 1992).

To mimic historic disturbance patterns, fire is being used 

as a management strategy in eastern deciduous forests. 

Despite the apparent benefits of prescribed burning, 

relatively little is known about how fire interacts 

with arthropod-dependent processes in eastern forest 

ecosystems. In this paper I review the evidence of direct 

and indirect interactions between forest arthropods and 

fire, and addresses the following questions: 1) are soiland litter-dwelling arthropod populations irreparably 

harmed by burning? 2) does prescription burning alter 

plant susceptibility to insect herbivores? 3) can fire 

be used as a management strategy to suppress forest 

arthropod pests?

Are soil- and litter-dwelling arthropods 

irreparably harmed by burning?

Relatively large, highly mobile arthropods are unlikely to 

be negatively impacted by prescribed fire. Oak savanna 

arthropod communities in the Upper Midwest were 

unaffected by burning (Siemann et al. 1997). In central 

Ohio, prescribed burning had no negative effect on large 

predatory carabid beetles (Smith and Horn 2000), or on 

scavenging scarab beetles (Stanton et al. 2000). Similarly, 

in south-central Kentucky, prescribed fire had little effect 

on ground-dwelling arthropod abundance, richness, and 

diversity, but arthropod community evenness increased 

in response to a single-burn disturbance (Coleman and 

Rieske, unpublished).

For less mobile arthropods functioning on the forest 

floor, the direct effects of fire include immediate 

mortality and habitat destruction. Loss of these faunal 

groups to fire could potentially disrupt essential 

ecosystem functions. One year following a prescribed 

fire in hardwood forests of eastern Kentucky there was 

a reduction in total dry mass of soil- and litter-dwelling 

invertebrates (Kalisz and Powell 2000). The loss was 

attributed primarily to direct mortality of Coleopteran 

larvae, which serve several critical roles on the forest 

floor as predators, herbivores, and detritivores (Dindal 

1990).

To examine the period of recovery needed to obtain 

pre-burn levels of arthropods following a prescribed fire 

regime using single- and multiple-burns, Coleman and 

Rieske (unpublished) monitored plots for two postburn growing seasons. Mites (Acari) and springtails 

(Collembola) dominate at the soil/litter interface, and 

populations were devastated by both fire regimes. Mite 

abundance did not recover over the course of the study. 

Arthropod abundance was affected by the prescribed 

burns, but arthropod diversity and richness were not. 

Litter arthropod evenness increased as a consequence 

of burning, probably due to reductions in the two 

dominant taxa.

In addition to the direct effects of habitat loss and 

mortality, burning reduces the available resource 

base, resulting in bottom-up regulation of soil/ litter 

community dynamics. Fire also may indirectly affect 

arthropod communities by changing plant species 

composition and foliar accessibility (Mitchell 1990), 

and by altering plant phenology. Changes in flowering 

phenology could affect insect pollinators, pollination 

rates, and subsequent seed set, though there are no 

studies addressing this.
1	 Fire 54 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Does prescription burning alter plant 

susceptibility to insect herbivores?

Fire suppresses vegetative competition and increases soil 

nutrients (Reich et al. 1990), and enhances seedling 

vigor (Adams and Rieske 2001), which may reduce the 

negative effects of herbivory. Fire also may indirectly 

affect arthropod communities by changing plant-species 

composition and foliar characteristics. Combustion of 

the litter layer provides an influx of nutrients to the 

forest floor, which can influence plant foliar chemistry 

through changes in nutrient availability and light 

intensity (Reich et al. 1990; Kruger and Reich 1997; 

Arthur et al. 1998), potentially altering herbivore feeding 

patterns (Rieske 2002; Rieske et al. 2002; Adams and 

Rieske 2003).

Fire-induced changes in foliar chemistry are not fully 

predictable and may be species-specific. Reich et al. 

(1990) and Kruger and Reich (1997) found enhanced, 

though transient, leaf nitrogen levels in northern red oak 

seedlings. Similarly, chestnut oak seedlings sampled in 

the post-burn growing season had higher foliar nitrogen 

and water content following a wildfire than seedlings 

sampled from unburned sites (Rieske 2002). By contrast, 

Kruger and Reich (1997) found a transient elevation 

in red oak seedling foliar carbohydrates following an 

early spring, low-intensity surface fire, whereas chestnut 

oak seedlings from burned sites had transient declines 

in foliar carbohydrates, and higher initial tannin levels 

(Rieske 2002). Although it is possible that chestnut oak 

and northern red oak seedlings respond differently to 

fire with respect to foliar carbon levels, the differences in 

seedling response may be attributed to the timing and 

intensity of the fires. Regardless, fire-induced changes 

may not be detectable, due to the buffering capacity of 

mature trees (Rieske et al. 2002), spotty fire coverage 

over irregular terrain, or to the relatively cool nature of 

some prescribed burns (Adams and Rieske 2003).

To assess the extent to which fire interacts with 

seedling herbivory, Adams and Rieske (2001) assessed 

mammalian and arthropod herbivore pressure on white 

oak seedling growth and vigor in burned forests in 

Kentucky. They found that herbivory is a measurable 

force impacting white oak seedlings but that prescribed 

fire did not affect herbivore pressure. They also found 

that the mammalian component of the herbivore 

complex had a greater impact on white oak seedling 

growth than the arthropod component.

Can fire be used as a management 

strategy to suppress forest arthropod 

pests?

Fire has been used extensively in agricultural, prairie, 

and rangeland ecosystems to manipulate vegetative 

composition and alter host plant quantity and quality, 

thereby directly or indirectly manipulating arthropod 

pest populations (Miller 1979). Successful suppression 

using fire requires that the pest be spatially and 

temporally vulnerable to fire-induced mortality (i.e., in 

the soil at the appropriate time), and that the fire itself 

does minimal damage to the standing crop of trees. 

Use of prescribed fire for pest suppression in managed 

forests has lagged behind that of other managed systems 

and has focused primarily on boreal forests (Mitchell 

1990; Brennan and Hermann 1994; McCullough et al. 

1998). The intricate link between fire and bark beetles 

as episodic disturbance agents in coniferous systems is 

now widely accepted, and recognition of this relationship 

gives land managers the opportunity to minimize 

harmful outcomes of these interactions.

Prescription burning has had limited use as a pest 

suppression strategy in deciduous forests of the 

Eastern United States (Brennan and Hermann 1994; 

McCullough et al. 1998). Historically, forest tracts in 

Massachusetts were burned to kill overwintering egg 

masses and clusters of feeding caterpillars as part of early 

attempts to eradicate the gypsy moth, Lymantria dispar, 

with mixed success (Doane and McManus 1981). More 

recently, prescribed fire was used to suppress populations 

of pear thrips, Taeniothrips inconsequens, in northern 

sugar maple stands. In addition to the direct mortality 

to overwintering thrips populations, fall burning seems 

to disrupt the phenological synchrony between emerging 

thrips and sugar maple bud expansion that is critical 

to the success of these insects (Brose and McCormick 

1992). Attempts to reduce acorn mortality caused by 

the acorn weevil, Conotrachelus posticatus, and associated 

acorn predators using prescribed fire in eastern oak 

forests have met with some success (Wright 1986; 

Roccardi et al. 2004; Rieske, unpublished). The highly 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 155

clustered spatial distribution of these seed feeders makes 

effective suppression and effective sampling problematic.

CONCLUSIONS

Although large, highly mobile arthropods appear to 

escape the negative effects of fire, the smaller, less mobile 

arthropods that are critical to decomposition processes 

are unable to escape. Arthropod abundance at the 

soil/litter interface can be severely reduced by burning, 

while effects on diversity, richness, and evenness are 

less marked. The use of repetitive prescribed fire to 

encourage regeneration and enhance stand development 

should be based on a schedule to allow appropriate time 

for leaf litter habitats to return to pre-burn conditions, 

and allow for resurgence of the arthropod community.

Fire-induced changes in oak foliar chemistry include 

increases in foliar nitrogen, carbohydrates, and 

defensive compounds, which can influence herbivore 

feeding patterns. Many of these changes are transient 

and not fully predictable; they may be species-specific 

and depend on the timing and intensity of the fire. 

Herbivory is a measurable force on oak seedlings, and 

mammalian herbivory has a greater impact on seedling 

vigor than arthropod herbivory.

Prescribed fire has limited use for pest suppression in 

deciduous forests of the Eastern United States. Pest 

vulnerability, both spatially and temporally, and the 

highly clustered spatial distribution of many pests 

makes effective suppression through prescription 

burning challenging and unpredictable. Although direct 

suppression of selected herbivores with prescribed fire 

is possible, prescription burning may be more viable 

as a means of enhancing forest health and redirecting 

succession, thereby reducing stand susceptibility to 

herbivore outbreaks.

Additional research is needed to fully understand the 

dynamics of insect-fire interactions in eastern oak forests. 

We lack a complete understanding of the effects of fire 

on the critical arthropod-driven processes in the soil/ 

litter interface. More work is needed on fire-induced 

changes in herbivore pressure and herbivore susceptibility 

of important forest tree species. 

ACKNOWLEdGMENTS

I thank Aaron Adams, Mary Arthur, Tom Coleman, 

and Heather Housman for their contributions, and the 

Daniel Boone National Forest for providing research 

sites. The comments and suggestions of three reviewers 

greatly strengthened this manuscript. This project was 

supported by the USDA Forest’s Service Southern 

Research Station and McIntire Stennis funds from 

the Kentucky Agricultural Experiment Station, and is 

published as Experiment Station Paper no. 06-08-009.

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1	 Fire 58 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

DOES FIRE AFFECT AMPHIBIANS AND REPTILES IN EASTERN U.S. 

OAK FORESTS?

Rochelle B. Renken1

Abstract.—Current information about the effect of fire on amphibians and reptiles in oak forests of the 

Eastern and Central United States is reviewed. Current data suggest that fire results in little direct mortality of 

amphibians and reptiles. Fire has no effect on overall amphibian abundance, diversity, and number of species 

in comparisons of burned and unburned plots, though salamander numbers tend to be greater in unburned 

plots. Current research also suggests that fire has no effect on reptile abundance, diversity, or number of 

species except in several studies in which lizard abundance was or tended to be greater in burned plots. The 

season of burn seems to make no difference in amphibian and reptile response. Although fire generally has 

little effect on amphibians and reptiles in oak forests, managers need to continue to consider the potential 

effect of fire on amphibians and reptiles associated with streams and forest pools, and on endangered 

threatened species and/or those of special concern. Managers can monitor the effects of fire on their own 

or with help from biologists, or can consult references to surmise how management affects these animals.

biomass of mice and shrews (Burton and Likens 1975). 

Reported densities for just the eastern red-backed 

salamander (Plethodon cinereus), one of the most 

common salamanders within oak forests, have reached 

11,452 per acre (Mathis 1991). Second, amphibians 

require moisture in the environment for breeding and 

respiration. Amphibians lay their eggs within water 

or in moist places (e.g., moss or logs) where eggs can 

remain moist or be covered by water. Amphibians also 

have permeable skin (Bury and others 2000) and are 

in a much greater danger of desiccation than birds 

or mammals. There are forest-dwelling salamanders 

(salamanders of the family Plethodontidae) that lack 

lungs and respire primarily through the skin. These 

species require moist environments for respiration. 

Third, amphibians and reptiles move relatively short 

distances (Bury and others 2000) such that they must 

deal with the changed conditions of their landscape and 

generally cannot escape or relocate to more favorable 

conditions (Szaro 1988). Amphibians and reptiles 

typically have small home ranges. For instance, eastern 

red-backed salamanders remain within an area of about 

15.5 square yards (Kleeberger and Werner 1983) of the 

forest floor within their lifetime; by contrast, the little 

brown skink (Scincella lateralis) has a home range of 62

square yards (Brooks 1967).

Fire within a forest can alter the environment for 

amphibians and reptiles in several ways. Forest floor litter 

and coarse woody debris, which are used by amphibians 

INTRODUCTION

Within the last 20 years, the effects of forest 

management on amphibians and reptiles has received 

increased attention. Typically, attention has focused 

on the effects of even-age and uneven-age forest 

management (see de Maynadier and Hunter 1995 for 

a review), but as fire has been gaining acceptance as a 

forest management tool, questions have been raised 

about the impact of fire (Russell and others 1999; 

Bury and others 2000; Pilliod and others 2003; Russell 

and others 2004). “Altered fire regime” was one of the 

reasons given for the problems experienced by 7 percent 

of 19 frog and toad species and about 17 percent of 49 

salamander species classified as “net extirpations” in the 

United States (Bradford 2005). Amphibians and reptiles 

are some of the least visible animals in the forest, yet 

with increasing focus on amphibian declines (Phillips 

1990; Wake 1991; Lannoo 2005), forest managers need 

to know the effects of their practices on these animals.

There are several reasons why amphibians and reptiles 

should be considered when making forest management 

decisions. First, they comprise a significant amount of 

biomass in forest systems. For instance, the biomass 

of salamanders alone within the Hubbard Brook 

Experimental Forest in New Hampshire was 2.6 

times that of birds and approximately equal to the 

1

Missouri Department of Conservation, 1110 S. College 

Ave., Columbia, MO 65201, 573-882-9909 ext. 3242, email: 

Rochelle.Renken@mdc.mo.gov.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 159

as a moist environment (Kleeberger and Werner 1983; 

Herbeck and Semlitsch 2000) and by reptiles as cover 

and perches (James and M’Closkey 2003), could be 

eliminated (Bury and others 2000; Pilliod and others 

2003; Vandermast and others 2004). Changes in canopy 

cover and ground level vegetation could result in a drier 

environment on the forest floor (Pilliod and others 

2003). The reduction or elimination of trees around 

temporary or permanent pools and streams could change 

the water temperature and chemistry, the amount of time 

water remains in pools, and the pattern of stream flow, 

thus changing the breeding environment for amphibians 

(Gresswell 1999; Skelly and others 1999; Pilliod and 

others 2003).

In the section that follows I summarize current 

information with respect to the effects of fire on forest 

amphibians and reptiles in oak forests of the Eastern 

and Central United States, and suggest issues that forest 

managers might to consider before using fire to obtain 

desired forest conditions.

CURRENT KNOWLEDGE ABOUT 

FIRE EFFECTS ON AMPHIBIANS AND 

REPTILES

Research on the effects of fire on amphibians and reptiles 

within oak forests of the Eastern and central United 

States has been limited, though studies to date indicate 

that fire results in little direct animal mortality, has little 

effect on amphibian abundance and diversity, and may 

result in increases in abundance for some reptile species. 

Palis (1995) in Indiana and Floyd and others (2002) in 

South Carolina did not find evidence of direct mortality 

of amphibians and reptiles due to fire. Direct mortality 

may not have occurred because amphibians tend to 

reside in moist environments, such as heavy litter and 

duff, that cannot sustain fire, or in underground refuges, 

such as small mammal tunnels (Gordon 1968), that 

would insulate them from fire (Russell and others 1999; 

Pilliod and others 2003). Reptiles also likely sought 

refuge in tunnels or under cover objects such as rocks, 

or climbed trees with the advance of fire. Although 

unburned sites tended to have more salamanders than 

sites that recently burned (the results between burned 

and unburned plots were not significantly different in 

statistical tests; Floyd and others 2002; Moseley and 

others 2003), investigators in Georgia (Moseley and 

others 2003), South Carolina (Floyd and others 2002), 

and Virginia (Keyser and others 2004) did not detect an 

effect of fire on overall amphibian abundance, diversity, 

or the number of species (Table 1). Nor did Ford and 

others (1999) observe an effect of fire on salamander 

numbers in North Carolina (Table 1). The only reported 

effect on amphibians was by Kirkland and others (1996), 

who observed more American toads (Bufo americanus) on 

burned than on unburned tracts in Pennsylvania (Table 

1). These authors did not have an explanation for this 

difference.

As for reptiles, some investigators have reported a 

greater abundance and diversity of lizards following 

burning (Table 1; Moseley and others 2003; Keyser and 

others 2004). These authors believed that the reduced 

litter, increased amount of bare ground, and removal 

of the midstory within burned forest resulted in more 

advantageous thermoregulatory conditions for lizards. 

However, others did not detect an impact of burning on 

lizard, snake, and turtle abundance and diversity, though 

lizards tended to be more abundant on burned plots 

(Table 1; Floyd and others 2002). Also, in preliminary 

analyses of more recent work, there were no effects of fire 

on amphibians in oak-hickory-poplar forests in South 

Carolina (K.R. Russell, University of Wisconsin-Stevens 

Point, pers. commun.) or on amphibian abundance 

(but perhaps on reptile abundance) in North Carolina 

oak-hickory forests (C. Greenberg, USDA Forest Service, 

pers. commun.). The season of burning also had no 

effect on amphibians and reptiles. Although spring and 

summer burns resulted in less forest shrub cover than 

winter burns, winter, spring, and summer burns did not 

differ in their effect on amphibian and reptile abundance 

(Keyser and others 2004). No investigators reported 

local disappearances of species or the appearance of new 

species on study plots following burning (Kirkland and 

others 1996; Ford and others 1999; Floyd and others 

2002; Moseley and others 2003).

In nearly all of the studies reporting the effects of fire 

on amphibians and reptiles in eastern oak forests, few 

discussed the intensity and type of fire (Table 1). The 

fires ranged from intense wildfires (Kirkland and others 

1996) and light prescribed burns (Floyd and others 
1	 Fire 60 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 Table 1.—Details of fires and amphibian and reptile responses to those fires in the eastern oak forests of the US. Location and reference Type of fire treatments Fire intensity Fuel consumption Patchiness of consumption Species e

valuated Animal r80esponse

Indiana (Palis 1995)

April burn; Prescribed burn set at 1615 hrs

Nighttime fire NAa 80 percent of 

1,000 acres burned

Amphibians and reptiles of that region

No direct mortality obser

ved; resulting 

amphibian and reptile numbers and assemblage appeared typical 

Pennsyl

vania 

(Kirkland and others 1996)

Burned in No

vember Charred tree 

trunks; killed many saplings; charred 

vegetation 

in ground and shrub layers

NA NA American toad More toads on burned 

than unburned sites

North Carolina (Ford and others 1999)

Burned in April High intensity 

prescribed fire

NA NA Woodland 

salamanders

No difference in salamander abundance detected between burned and unburned sites

South Carolina Piedmont (Floyd and others 

200

2)

First round of fires in February - March 1999; strip head fires set 15 to 30 feet apart; second round of fires in April 

2000; 

strip head fires set 10 to 

20 feet apart; 

First round average 

flame length was >1 foot; second round average flame length < 1foot

NA NA Total herpetofauna, 

frogs, toads, salamanders, newts, lizards, turtles, snakes

No difference in abundance, di

versity, 

and e

veness in total 

herpetofauna or by taxonomic groups
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 161

Georgia upper Coastal Plain (Moseley and others 

2003)

Winter burns; stands had been burned every 

2-3 years in the 

pre

vious 9 years

Low intensity More litter in 

unburned; more bare ground in burned; no difference in coarse woody debris volume between burned and unburned

NA Total herpetofauna; 

amphibians, anurans, salamanders, reptiles

No difference in o

verall amphibian and 

reptile, salamander, or anuran abundance, richness, and di

versity 

between burned and unburned; reptile abundance and di

versity were greater 

in burned than unburned sites

Virginia Piedmont (Keyser and others 

2004)

Seasonal prescribed burns (winter, spring, summer) applied to first-stage shelterwood-har

vested 

stands (to 48 ft2 basal area per acre) cut 3 to 5 years prior to burns

Average fire 

temperature 39 inches abo

ve the 

ground was 5

26 

°F in winter burn, 648 °F in spring burn, and 485 °F in summer burn

All fires reduced fine, medium, coarse fuel loads (fine fuels were eliminated; coarse fuels were partially reduced), yet duff layer remained intact

Spring and summer burns caused a shift from shrub dominated to herbaceous dominated understory

All amphibians and reptiles, especially eastern red-backed salamanders, American toads, northern fence lizards (Sceloporus undulatus), little brown skinks, and southeastern fi

velined skinks (Eumeces 

inexpectatus) 

No difference in abundance of salamanders, toads, and all amphibians between controls and different seasonal burns; lizard abundance was greater in burned than unburned stands

a Not Available

Table 1.—continued.
1	 Fire 62 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

2002; Moseley and others 2003) to high-intensity 

prescribed burns (Ford and others 1999; Keyser and 

others 2004). The effects of fire on amphibians and 

reptiles generally were similar among these studies.

Researchers may not have detected an effect of fire on 

amphibians and many reptiles due to short-term changes 

in the physical features important to these animals or 

the effects were not significant. Unless conditions are 

extremely dry, some coarse woody debris remains after 

a fire (Palis 1995; Van Lear and Harlow 2000; Moseley 

and others 2003; Trammell and others 2004) and can 

serve as cover for both amphibians and reptiles. Fire 

causes a temporary reduction in ground flora cover and 

litter depending on the frequency and intensity of fire, 

but ground flora cover rebounds quickly, sometimes 

within the next growing season (Kirkland and others 

1996; Vandermast and others 2004). Not all litter or 

duff is consumed (Fig. 1; Kirkland and others 1996; 

Floyd and others 2002; Keyser and others 2004; 

Trammell and others 2004) and burns may create a 

mosaic of vegetation on the forest floor (Ford and 

others 1999). Litter seems to accumulate to unburned 

plot levels within 3 years after fire (Gagan 2002). Also, 

unless the objective of burning is to dramatically reduce 

the density of trees and greatly open the canopy, trees 

remaining following infrequent fires provide shade to 

reduce the amount of temperature variation at the forest 

floor, and the canopy helps soil and any remaining litter 

retain moisture (Ford and others 1999). Moisture within 

soil, litter, and coarse woody debris allows salamanders 

to remain active at the forest floor surface. In dry 

environments, amphibians, particularly salamanders, 

retreat underground and remain there until conditions 

on the forest floor are more suitable (Heatwole 1960; 

Heatwole 1962; Jaeger 1980; Feder 1983). For lizard 

species that experience an increase in abundance with 

fire, the temporary benefits of more bare ground and 

more basking sites probably boost survival rates (Parker 

1994). However, populations probably return to preburn levels as post-fire forest conditions return to 

conditions of unburned forest.

OTHER CONSIDERATIONS 

CONCERNING THE EFFECTS OF FIRE

Do we need to be aware of other considerations 

concerning the use of fire as a tool and its effect on 

amphibians and reptiles? One is the possible indirect 

effects of fire on streams and stream-associated 

amphibians. There have been no studies on the indirect 

effects of fire on streams in eastern oak forests, but it 

is possible that streams could be affected by increased 

rates of sedimentation, increases in temperature due to 

reduced canopy cover, increases in amount of woody 

debris into the stream, which could change the flow 

and course of the stream, and increased concentrations 

of chemicals from the leaching of ash and the 

diffusion of smoke (Gresswell 1999; Pilliod and others 

2003). Increased rates of sedimentation, increased 

concentrations of chemicals such as potassium and 

Figure 1.—Results of a prescribed burn 

where not all the litter and woody debris 

was eliminated.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 163

nitrogen, and increases in woody debris may change 

stream productivity and habitats, thus changing the flow 

and quality of streams. Stream-associated amphibians 

may be sensitive to such changes (Bury and others 2000; 

Pilliod and others 2003), which also can affect the foods 

and reproduction of stream-side dwelling amphibians, 

such as the northern dusky salamander (Demognathus 

fuscus). A reduced canopy cover might result in increased 

water temperature, which also could affect foods (e.g., 

macroinvertebrates) and reproduction of these animals 

(Pilliod and others 2003).

Another consideration is the direct effect of fire on 

vegetation structure around amphibian breeding ponds. 

Frequent fires open the canopy around pools and the 

increased light can change amphibian use of the pools. 

Depending on how isolated the pool is from other pools 

and the amount of time water is in the basin in a year, 

amphibian species more characteristic of open pools may 

start using formerly closed-canopy pools. Amphibian 

species such as spring peepers (Pseudacris crucifer) prefer 

open habitats and more open canopies around breeding 

pools (Skelly and others 1999; Halverson and others 

2003), and may invade formerly closed-canopy pools 

if other spring peeper populations are nearby. Pools in 

closed canopies typically have lower oxygen levels and 

water temperatures, and shorter periods when water is in 

the basin than open-canopy pools of the same category 

(Skelly and others 1999, 2002; Pilliod and others 2003). 

The reason for this is that trees shade the water and use 

the water within the basin (Skelly and others 1999). 

Lower water temperatures reduce tadpole growth rates 

for some amphibian species (Skelly and others 2002; 

Halverson and others 2003). Reduced growth rates and 

faster drying pools in closed canopies make it difficult 

for tadpoles to develop into terrestrial-living juveniles 

before the pools dry. In a study examining amphibians 

using ponds in Michigan between 1988 and 1992, Skelly 

and others (1999) noted that tree growth and maturation 

around pools that were open-canopy pools from 1967 

to 1974 resulted in fewer amphibian species using the 

pools. The composition of the amphibian community 

was less diverse and species that preferred more open 

habitats stopped using the closed-canopy pools. The 

remaining amphibians were considered forest species and 

generally tolerated of a wider range of canopy conditions 

than open-habitat species.

A third consideration is the use of fire for forest 

management in the presence of endangered or 

threatened species and/or those of special concern. 

Although direct mortality does not seem to be an issue, 

managers may want to be aware of these species’ habitat 

needs and seasonal and daily activity patterns so that fire 

does not result in direct mortality (e.g., burning outside 

the months of the year when that species is active), and 

that required habitat remains or is improved following 

burning.

A fourth consideration is that while current information 

suggests that infrequent fires in eastern oak forests result 

in little direct mortality or in insignificant impacts to 

amphibians and limited changes to reptile communities, 

future data may point to impacts that have not yet been 

measured. I found few published studies on the effects 

of fire on amphibians and reptiles in eastern oak forests, 

yet I am aware that several other studies that are in 

progress or that recently were completed (C. Greenberg, 

USDA Forest Service, pers. commun.; D. Miles, Ohio 

University, pers. commun. part of the National Fire 

and Fire Surrogate Study; K. Russell, University of 

Wisconsin-Stevens Point, pers. commun.).

CONCLUSIONS

Currently available information indicates that fire 

has little effect on amphibians and reptiles in oak 

forests of the Eastern and Central United States. Some 

researchers have noted fewer salamanders in burned 

areas than unburned areas, though significant changes in 

amphibian abundance, diversity, and number of species 

have not been detected. Researchers have also reported 

that fire generally does not affect reptile abundance 

and diversity but that there is evidence that lizard 

abundance increases following burning. The lack of 

significant effects likely is due in part to the incomplete 

elimination of moisture-holding litter, duff, and coarse 

woody debris, the continued canopy cover provided 

by remaining trees, and the quick regrowth of ground 

vegetation (Palis 1995; Moseley and others 2003; Keyser 

and others 2004).
1	 Fire 64 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Forest managers must consider many factors when 

contemplating management goals on properties under 

their care. In an ideal situation, a manager would 

monitor the impacts of management actions on 

amphibians and reptiles along with such monitoring 

as necessary to determine whether the desired forest 

conditions are obtained. Few managers have the luxury 

of time and money to monitor amphibian and reptile 

communities. The next best option is to work with 

a biologist who can monitor amphibian and reptile 

responses or provide information to help guide decision 

making. If this option is not available, the manager can 

consult general references related to the biology and 

habitat needs of nontarget animals. Examples include 

“A field guide to reptiles and amphibians of eastern and 

central North America” (Conant and Collins 1991), 

“A guide to amphibians and reptiles” (Tyning 1990), 

and “Salamanders of the United States and Canada” 

(Petranka 1998). “Amphibian declines: the conservation 

status of United States species” (Lannoo 2005) provides 

information on the life history, natural history, and 

conservation status of all amphibians in the United 

States. Other helpful references include state or regional 

publications such as “The amphibians and reptiles of 

Missouri” (Johnson 2000)

ACKNOWLEDGMENTS

I thank the Missouri Department of Conservation 

for providing financial and logistical support and the 

researchers and technicians who have worked on this 

topic and communicated their results in publications. 

I also thank M. Anderson, M. Huffman, J. Tuttle, and 

two anonymous reviewers for reviewing early drafts of 

this paper.

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Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 167

Conservation implications for neotropical migratory

and game birds in oak-hardwood stands managed with

shelterwood harvests and prescribed fire

J. Drew Lanham1†, Patrick H. Brose2 and Patrick D. Keyser3

Abstract.—Prescribed fire in conjunction with shelterwood cutting is a novel way to regenerate oakdominated stands on certain upland sites while minimizing the intrusion of hardwoods. We describe 

three options (complete or partial canopy retention, postharvest prescribed burning, and complete canopy 

removal) within a shelterwood-prescribed fire regime that will create two-age stands that are likely to 

harbor a diverse mixture of mature forest and early successional birds; parklike woodlands with open 

woodland species; or early successional habitats with shrubland species. This system is a viable option for 

managing both avian and timber resources where oak-dominated stands on upland sites are the desired goal.

more efficient means of regenerating oaks in shelterwood 

systems. Keyser et al. (1996) burned two oak-dominated 

shelterwood stands after an initial harvest and found that 

regeneration of yellow-poplar, red maple, and sweetgum 

(Liquidambar styraciflua L.) was reduced by as much as 

90 percent while oak reproduction was reduced only by 

11 percent. Followup investigations of fire effects in oakdominated shelterwood stands reported similar results 

in high fire resistance among oak and low tolerance for 

burning among less desirable hardwood competitors. 

This illustrates the vital role of postharvest burning 

coupled with growing-season burns in creating an oakdominated seedling cohort (Brose and Van Lear 1998a,b; 

Brose et al. 1999; Brose and Van Lear 1999; Van Lear 

and Brose 1999).

We suggest that this technique may be applicable 

elsewhere in upland oak-dominated stands (oak-hickory) 

of Eastern North America (Ward and Gluck 1999). 

Application of this technique (Fig. 1a-h) entails threesteps. First, an initial harvest leaves 50 to 60 dominant 

oaks per ha (11 to 12 m2

 of basal area/ha). The remnant 

stand should contain the best oak stock to encourage 

a vigorous regeneration cohort. The stand is then left 

undisturbed for 3 to 5 years to allow the development 

of a regeneration layer. After 3 to 5 years, a hot (flame 

length > 1.0 m) growing-season fire is applied to the 

stand, resulting in an oak-dominated regeneration 

cohort.

Although this technique was originally implemented 

to improve the viability of oak regeneration and the 

Introduction

In Eastern North America, shelterwood silviculture is a 

common technique for managing oak-dominated stands 

on upland sites (Sander et al. 1983). Partial harvests 

reduce the dense shade that suppresses root development 

of existing oak regeneration (Loftis 1990) and helps retard 

the rapid height growth of less-desirable species such as 

yellow poplar (Liriodendron tulipifera L.) and red maple 

(Acer rubrum L.). Additionally, soil disturbance from 

the harvesting operations prepares seedbeds for acorns 

produced by the retention stand, thereby encouraging oak 

seedling establishment (Cook et al. 1998).

On some upland sites, preharvest treatments such as 

herbicide application (Loftis 1990; Lorimer et al. 1994), 

low-intensity burning (Barnes and Van Lear 1998; 

McGill et al. 1999), tree sheltering (Potter 1988), and 

implantation of high-quality nursery stock (Bowersox 

1993; Gordon et al. 1995; Schlarbaum et al. 1997) may 

precede the initial shelterwood harvest to encourage oak 

regeneration. However, these preharvest treatments are 

expensive and often ineffective and must precede the 

initial shelterwood cut by 5 to 15 years to allow sufficient 

root development of the oak regeneration.

The constraints of time and money that often affect 

private owners of small- to medium acreage require a 

1

Clemson University, Department of Forestry and Natural 

Resources, 261 Lehotsky Hall, Clemson, SC 29634; 2USDA 

Forest Service, Northern Research Station, Irvine PA 16329; 3

Forestry Division, MeadWestvaco Corporation, P.O. Box 577, 

Rupert, WV 25984; †Corresponding author, 864-656-7294, 

email: lanhamj@clemson.edu.
1	 Fire 68 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

production of hardwood timber in uplands, the 

conservation of biodiversity also is a goal of many 

forest management activities. Birds comprise significant 

ecological components of eastern forest systems. 

Songbird population declines in many forests of the 

Eastern United States have focused attention on the 

effects of various silvicultural practices on conservation 

efforts. Such efforts have been particularly focused on 

many species of neotropical migrants (Askins et al. 

1990). Game birds have long been a focus of forest 

management because of their economic and recreational 

importance. While numerous investigators have 

reported the effects of various silvicultural treatments 

on game and nongame birds, there is a dearth of 

information addressing the effects (real or potential) of 

prescribed fire on avifauna in hardwood systems. We 

discuss how various management options implemented 

in oak­- shelterwood burn systems can influence the 

composition of avian communities. These potential 

influences are related to the vegetative structure and 

composition that result from the three oak-shelterwood 

burn options and from inferences drawn from other 

bird habitat studies conducted in other hardwood or 

pine- hardwood systems where silvicultural treatments 

have created habitat conditions similar to those expected 

in oak-shelterwood burns. To emphasize the potential 

importance of these treatments from a bird conservation 

perspective, we have used Bird Conservation Region 

(BCR) breeding scores (RCS-b) from Region 24 

– Central Hardwoods, Region 28 – Appalachian 

Mountains, and Region 29 – Piedmont (Partners in 

Flight Species Assessment database: ww.rmbo.org/

pif/scores/scores.html) as indicators of management 

priority.

Conservation Implications for Avian 

Communities

Canopy Retention

Canopy retention treatments provide two-aged stands 

twice during the shelterwood burn regime: 1) when 

“undesirable” hardwoods dominate the advanceregeneration cohort, and 2) after a satisfactory cohort of 

vigorous, advance oak- regeneration is achieved when 

a portion or all of the residual overstory trees may be 

retained for at least half of the next rotation.

A widely observed trend of bird -habitat relationship 

studies is the positive association between floristic 

structural diversity and bird-species diversity. Retention 

of some overstory during shelterwood treatments 

may provide sufficient canopy habitat and vertical 

structure for some mature forest canopy bird species 

to use partially harvested stands (Dickson et al. 1995). 

Relative to the composition of a mature forest songbird 

community, canopy retention treatments would be the 

least intensive and probably most similar to an unevenaged mature forest. Dickson et al. (1995) concluded that 

the retention of a residual canopy (less than 50 percent) 

for several years after an initial harvest can provide 

habitats for some mature forest birds that otherwise 

would not inhabit stands managed using traditional 

even-age management techniques.

Wood and Nichols (1995) found that two-aged stands 

in West Virginia contained a greater density, richness, 

evenness, and overall diversity of breeding birds than 

early successional or mature stands. The total density 

for all neotropical migrants also was highest in the 

two-aged stands. Densities of forest-interior species 

did not differ statistically among clearcut, mature, and 

two-aged stands. The two-aged stands had densities of 

interior-edge species equal to or greater than the other 

two treatments. The co-occurrence of mature forest and 

early successional species within the same areas indicated 

that two-aged stands provided habitats for both mature 

and early successional species. Mature forest species 

reported in The Wood and Nichols study included veery 

(scientific names for all neotropical migrant and game 

bird species are in Table 1), American redstart, and 

scarlet tanager. Early successional species recorded in 

the same stands included chestnut-sided warbler, indigo 

bunting, and eastern towhee.

In the Missouri Ozarks, Annard and Thompson (1997) 

reported higher species richness for breeding birds 

in stands treated by the shelterwood method than in 

clearcuts, group selection, single-tree selection, or uncut 

stands. Species richness was higher in shelterwoods than 

in unharvested or uneven-aged stands. As in the Wood 

and Nichols study, these differences were attributed to 

the presence of both early seral stage and mature forest 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 169

Table 1.—Oak shelterwood burn occupancy potential of select neotropical migrant and game bird speciesa

Species Option 1 Option 2 Option 3

wild turkey (Meleagris gallopavo) X X X

northern bobwhite (Colinus virginianus) X X

ruffed grouse (Bonassa umbellus) X X X

yellow-billed cuckoo (Coccyzus americanus) X

black-billed cuckoo (C. erythropthalmus) X

whip-poor-will (Caprimulgus vociferus) X

Chuck-will’s-widow (C. carolinensis) X

ruby-thr. hummingbird (Archilocus colubris) X X

Acadian flycatcher (Empidonax virescens) X

least flycatcher (E. minimus) X

willow flycatcher (E.alnorum)

eastern kingbird (Tyrannus tyrannus) X

great-crested flycatcher (Myiarchus crinitus) X

eastern wood-pewee (Contupus virens) X

gray catbird (Dumatella carolinensis) X X

wood thrush (Hylocichla mustelina) X

veery (Catharus fuscescens) X

blue-gray gnatcatcher (Pilioptila caerulea) X

red-eyed vireo (Vireo olivaceous) X

yellow-throated vireo (V. flavifrons X

warbling vireo (V. gilvus) X

white-eyed vireo (Vireo griseus) X

Blackburnian warbler (Dendroica fusca) X

black-throated blue warbler (D. caerulescens) X

black-throated green warbler (D. virens) X

cerulean warbler (D. cerulea) X X

chestnut-sided warbler (D. pensylvanica) X X

yellow-throated warbler (D. dominica) X

prairie warbler (D. discolor) X

yellow-warbler (D.petichia) X

blue-winged warbler (Vermivora pinus) X X

golden-winged warbler (V. chrysoptera) X X

American redstart (Setophaga ruticilla) X

black-and-white warbler (Mniotilta varia) X X

common yellowthroat (Geothlypis trichas) X

hooded warbler (Wilsonia citrina) X

Kentucky warbler (Oporonis formosus) X

northern parula (Parula americana) X

ovenbird (Seiurus aurocapillus) X

Continued
1	 Fire 70 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Louisiana waterthrush (S. motacilla) X

worm-eating warbler (Helmitheros vermivorus) X

yellow-breasted chat (Icteria virens) X

orchard oriole (Icterus spurius) X X

Baltimore oriole (I. galbula) X X

scarlet tanager (Piranga olivacea) X

summer tanager (P. rubra) X

indigo bunting (Passerina cyanea) X X X

blue grosbeak (Guiraca caerulea) X X

a

Bird habitat associations are inferred from associations in similar types and conditions of habitat that would be created by 

the options described in this paper. Bird habitat associations are derived from Hamel et al. (1982) and denote associations 

with seral stages in oak-hickory habitats as follows: Option 1 = sapling poletimber-sawtimber; Option 2 = grass-forb, 

sawtimber; Option 3 = seedling-sapling.

Table 1.—Continued.

Species Option 1 Option 2 Option 3

birds, including blue-winged warbler, prairie warbler, 

red-eyed vireo, worm-eating warbler, and Acadian 

flycatcher.

Nesting success must be considered in conjunction with 

measures of density and diversity of breeding birds. 

Wood and Nichols (1995) reported no differences in 

nest success among treatments in West Virginia. Nest 

parasitism by brown-headed cowbirds (Molothrus ater) 

was not a major factor in their study; only 8 of 246 

nests were parasitized and there were no differences in 

the number of cowbirds among treatments. Annard 

and Thompson (1997) and Welsh and Healy (1993) 

reported similar results in Missouri and New Hampshire, 

respectively. However, one must remain aware, that 

patterns of predation and parasitism may vary depending 

on the landscape context. Overall, the impact of 

cowbirds and predators in extensively forested systems 

tends to be lower than that in agricultural and suburban 

landscapes (Wilcove 1985)

The findings of all of these studies are consistent with 

an earlier study by Crawford et al. (1981). Although 

they did not study shelterwood systems, the findings 

of Crawford et al. nonetheless are relevant because 

they concluded that timber management strategies 

altered bird communities in relationship to the degree 

of stand disturbance. They predicted that partial 

harvests would provide sufficient canopy cover to buffer 

complete species turnover from mature forest to early 

successional species observed in clearcut forests. They 

further surmised that partial cuts would return more 

quickly to site conditions conducive to mature bird 

species than even-age treatments. These findings have 

been corroborated by other studies that have shown 

that although populations of some forest-interior 

songbirds may be reduced relative to an undisturbed 

stand due to habitat alteration, increased nest predation, 

and parasitism (Webb et al. 1977; Wood and Nichols 

1995), these species generally are not eliminated entirely 

and population recovery may occur rapidly as the new 

forest matures (Conner and Adkisson 1975; Askins and 

Philbrick 1987).

Canopy disturbance may benefit forest-interior bird 

species that have declined in some regions. Some birds 

that use early-successional gaps within mature forests 

may decline in areas where disturbances do not produce 

the regenerating ground-layer and shrub vegetation they 

prefer (Franzreb and Rosenberg 1997). Shelterwood 

harvesting would stimulate the growth of low vegetative 

cover. In West Virginia, Wood and Nichols (1995) 

found that Kentucky warblers, wood thrushes, American 

redstarts, and black-and-white warblers were 2 to 3 

times more abundant in two-aged stands than in uncut 

controls. The abundance of these species in the shrubbier 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 171

two-age stands lends credence to the shelterwood-shrub 

layer hypothesis.

The retention of the best, 11 to 12 m2

 of oak basal area/

ha (50 to 60 dominant oaks/ha) in shelterwood stands 

also provides reliable acorn sources (Healy 1997). Acorns 

are one of the most important wildlife foods as they 

are eaten by more than 200 wildlife species throughout 

North America (Martin et al. 1951). Among these are 

a multitude of avian species (Martin et. al 1951; Beck 

1993). Corvids, e.g., blue jays (Cyanocitta cristata) and 

American crows (Corvus brachyrhynchos), are voracious 

acorn predators and bear a large responsibility for the 

regeneration of oak stands through their caching activity. 

Although not a neotropical migrant or game species, 

red-headed woodpeckers (Melanerpes erythrocephalus) are 

short-distance migrants that are likely to benefit from 

overstory retention management activities. They also 

receive attention as species of regional importance from 

Partners in Flight (Panjabi et al. 2005). Several species 

of upland game birds including wild turkeys and ruffed 

grouse consume acorns (Martin et al. 1951; Dickson 

2001), making canopy retention stands potentially 

important foraging habitat for both species.

On the basis of the floristic structure of stands expected 

after canopy retention treatments, Table 1 lists the 

diverse array of neotropical migratory and game bird 

species that are likely to occur in the diverse two-age 

structure of these areas. Conservation priority scores 

and management priorities for species occurring under 

this option are shown in Table 2. Note that the cerulean 

warbler, a species likely to benefit from the mature 

canopy retained in this silvicultural scenario (in mesic 

types), is characterized as a species with “Immediate 

Management” conservation priorities in both the 

Central Hardwoods and Appalachian Mountain Bird 

Conservation Regions BCR (Panjabi et al. 2005). 

Illustrating the diverse avian conservation potential for 

this scenario, golden-winged warblers and ruffed grouse 

also are Immediate Management priorities (Panjabi et al. 

2005). In the Piedmont, conservation priorities are even 

higher as ruffed grouse are characterized as a species in 

need of “Critical Recovery”. Ruffed grouse might occupy 

the complex woody/herbaceous understory resulting 

from the overstory retention option in the Central 

Hardwoods, Appalachian, and Piedmont BCR while 

golden-winged warblers should occupy similar habitats 

in the Appalachian mountains.

Understory Suppression Using Prescribed 

Burning

The second option in the oak-shelterwood burn scheme 

is the use of periodic prescribed fire in shelterwood 

stands. Among the three options discussed here, this 

method is likely to be intermediate in its effects on the 

songbird community. Ultimately, the shift in species 

composition will vary depending on the vegetative 

structure resulting from the season, intensity, and 

frequency of the prescribed fire. Dormant-season 

(winter) burns will produce low-growing, sprouting 

regeneration of shrub and trees and stimulate the 

production of soft mast (Stransky and Roese 1984). 

These responses may provide forage, cover and arthropod 

prey for many game and nongame birds (Dickson 1981, 

2001).

Repeat dormant-season burning promotes an increased 

abundance of oak regeneration. Oak regeneration is 

limited with additional fires and released at intervals 

by withholding burning treatments, creating patchy 

stands in different successional stages. Dickson (1981) 

surmised that in southern pine and pine-hardwood 

forests, a patchwork of different successional stages 

within a stand (or across a landscape) could enhance 

bird diversity and abundance. This patchwork obviously 

would be dependent not only on the frequency and 

intensity of fires but also on the size, topography, and 

site capability of the area burned. In stands managed 

with dormant-season fires that allow the proliferation 

of hardwood shrubs and trees underneath an open 

canopy, bird communities are likely to consist of a large 

proportion of shrub nesting, e.g., white-eyed vireo, and 

midstory species, e.g., wood thrush, along with species 

more characteristic of open canopy forests, eg., yellowbilled cuckoo and blue-gray gnatcatcher. More so than 

other burning treatments, dormant-season fires in oakshelterwoods are likely to produce bird communities 

more similar to two-aged canopy retention stands.

Annual or biennial prescribed burning during the 

growing season should create open hardwood woodlands 
1	 Fire 72 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Table 2.—Conservation scores and management prioritiesa for selected avian species of 

regional importance potentially occupying oak-shelterwood burn habitats in central 

hardwoods (CH), Appalachian mountain (AM), and piedmont (PM) bird conservation 

regions.

Species CH AM PM

northern bobwhite 16 MA 15 IM 16 IM

ruffed grouse 15 IM 16 IM 14 CR

yellow-billed cuckoo 15 MA 14 MA

black-billed cuckoo 17 MA 14 MA

whip-poor-will 17 MA 16 MA 18 MA

Chuck-will’s-widow 14 MA 15 MA

Acadian flycatcher 16 PR 17 MA

willow flycatcher 12 PR 11 PR 9 PR

eastern kingbird 15 MA 14 MA

eastern wood-pewee 15 MA 15 MA 15 MA

wood thrush 16 MA 16MA 16 MA

blue-gray gnatcatcher 14 MA

yellow-throated vireo 16 PR 17 MA 14 PR

white-eyed vireo 15 MA

Blackburnian warbler 14 MA

cerulean warbler 19 IM 21 IM 16 MA

prairie warbler 18 MA 18 MA 18 MA

yellow-throated warbler 15 PR 16 PR

blue-winged warbler 19 MA 17 PR 16 MA

golden-winged warbler 21 IM

black-and-white warbler 16 MA

hooded warbler 15 PR

Kentucky warbler 18 MA 19 MA 15 PR

Louisiana waterthrush 15 PR 18 MA

worm-eating warbler 18 MA 18 MA 13 PR

yellow-breasted chat 16 MA 15 MA

orchard oriole 17 MA

Baltimore oriole 14 MA

scarlet tanager 14 PR

summer tanager 6 PR 16 MA

indigo bunting 14 PR 14 PR 12 PR

blue grosbeak 14 PR

Average conservation score 18.22 16.24 14.6

Critical recovery 0 0 1

Immediate management 2 4 1

a

Management actions in decreasing priority: CR= Critical Recovery; IM = Immediate Management; 

MA = Management Attention; PR = Planning and Responsibility (Panjabi et al. 2005)
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 173

and savannas by gradually eliminating much hardwood 

shrub and tree regeneration while stimulating production 

of ground-level herbaceous vegetation (Thor and 

Nichols 1973). Oak woodland and savanna habitats 

were described as common landscape features by early 

explorers and settlers who observed the extensive use 

of fire by indigenous Americans (Pyne 1982; Buckner 

1983; Van Lear and Waldrop 1989). However, over time 

oak savannas and woodlands and some of the wildlife 

species associated with them have become rare. The 

restoration of hardwood savannas and open woodlands 

probably would shift bird guilds from mature forestinterior species to canopy and midstory dwelling, open 

woodland and grove species, e.g., great-crested flycatcher, 

eastern wood-pewee, orchard oriole and summer tanager. 

Here, even more so than in overstory retention stands, 

the red-headed woodpecker is likely to benefit from this 

management option. Such open habitats may also likely 

to attractive to wild turkey for a number of life requisites 

including foraging, resting, and brood rearing.

Although growing-season fires might benefit some bird 

species, others could be negatively impacted by burns 

initiated so late that nesting and other breeding activities 

are disrupted. Therefore, spring burning should be 

prescribed judiciously as early as possible in the season to 

minimize direct impacts on nesting or breeding birds.

Fire intensity (hot versus cool) also affects vegetative 

structure and, therefore, avian community composition. 

Studying the effects of fire intensity on bird communities 

in Alabama Piedmont pine-hardwoods, Stribling 

and Barron (1995) found a greater abundance and 

diversity of birds in less intensively burned stands with 

canopy, shrub, and cavity nesters the most abundant. 

Canopy, shrub, and bark-feeding species also were more 

abundant in cool burn sites than in untreated stands. 

These differences were attributed to the patchiness of 

the vegetative structure in these areas. This same study 

reported a higher abundance of ground-foraging and 

ground-nesting songbirds in stands burned with a hot, 

growing-season fire than in those burned by cooler 

growing-season fires. They surmised that the observed 

responses of terrestrially associated species could have 

occurred because of litter removal that may have 

provided better foraging and nesting habitat.

Some residual canopy trees such as maples (Acer spp.) 

and tulip-poplars and those with slash accumulations at 

their bases are susceptible to fire-kill or damage (Brose 

and Van Lear 1999a). These snags are important foraging 

sites for woodpeckers and other bark-gleaning species. 

Snags also provide perching/hawking sites and roosting/

nesting habitats. Larger snags provide nesting habitats 

for both primary cavity excavators (woodpeckers) and 

secondary cavity nesters, including neotropical migrants 

such as the great-crested flycatcher (Lanham and Guynn 

1996). In addition to the valuable function provided by 

snags, downed logs and other coarse woody debris, e.g., 

treetops, fallen limbs, provide habitat for forest floordwelling arthropods, herpetofauna, and small mammals 

(Hanula 1996; Loeb 1996, Whiles and Grubaugh 1996). 

These provide food resources for songbirds and larger bird 

species, e.g., raptors, wild turkey. Larger logs remaining 

on the forest floor also may provide drumming substrate 

for ruffed grouse (Dickson 2001). Because the effects 

of fire in forested stands can have such varied effects, a 

variety of bird species is possible based on fire frequency 

and intensity and various site characteristics. Because 

most natural resource mangers and private landowners are 

primarily concerned with the production of open, oakdominated woodland, Table 1 lists species likely to occur 

in understory (growing-season) burned treatments that 

result in park-like, oak woodlands. Conservation priority 

scores for species occurring under this option are shown 

in Table 2. Of note within the context of an option 

producing woodland species, highest conservation scores 

and management priorities are listed for the cerulean 

warbler, which may benefit from the park-like conditions 

should they be implemented in mesic oak/hardwoods 

preferred by the species in the Central Hardwoods and 

Appalachian Mountain BCR. Ruffed Grouse retain 

an Immediate Management priority for this option in 

Central Hardwoods and Appalachian mountain habitats. 

In the Piedmont, this priority increases to Critical 

Recovery status

Overstory Removal

The third option of harvesting all of the residual 

overstory trees creates early successional hardwood 

habitats. These stands will undergo a dramatic turnover 

in avian composition with species such as indigo 

buntings and field sparrows occurring in regenerating 
1	 Fire 74 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

hardwood stands during the grass-forb and seedlingsapling stages (Evans 1978). In subsequent years, as 

vertical structure within a regenerating stand changes 

with the growth of shrubs and saplings, avian diversity 

and abundance may surpass those of mature stands 

(Conner and Adkisson 1975; Thompson and Fritzell 

1990; Thompson et al. 1992). In many eastern uplands, 

regenerating seedling-sapling hardwood habitats are 

preferred by shrub-scrub species such as prairie warblers, 

yellow-breasted chats, and chestnut-sided warblers. At 

this juncture in succession, northern bobwhite quail 

may use woody or brushy hardwood habitats that are 

interspersed with other cover types, e.g., pine forests, 

fallow fields, as escape, resting, and roosting cover in the 

nonbreeding season (Dickson 2001). Early successional 

hardwood stands may be used by wild turkey for nesting 

and brood-rearing habitat (Speake et al.1975). Ruffed 

grouse are especially dependent on dense thickets of 

hardwoods for drumming and brood rearing. Evenage systems including shelterwood regimes are cited 

by Dickson (2001) as a preferred method for creating 

suitable habitat for the species.

As regenerating stands mature to form closed canopy 

sapling-poletimber stands, species richness and 

abundance frequently decrease to levels below younger 

shrubland and older mature forest habitats (Conner and 

Adkisson 1975). However, some forest-interior songbirds 

such as black-and-white warblers and wood thrushes 

will begin using pole stands at this stage (Conner and 

Adkisson 1975; Mauer et al. 1981; Askins and Philbrick 

1987). These stands continue to hold high suitability for 

ruffed grouse. (Gullion 1984). Table 1 lists bird species 

typical of regenerating, early successional hardwood 

stands. Conservation priority scores for species occurring 

under this option are shown in Table 2. Given the 

early successional conditions that are reproduced in 

this option, highest conservation priorities are given to 

the ruffed grouse (Central Hardwoods, Appalachian 

Mountains, Piedmont), northern bobwhite (Appalachian 

Mountains, Piedmont), and golden winged warbler 

(Appalachian Mountains).

Discussion

The hardwood forests of Eastern North America are 

one of the largest, broad-leaved, deciduous ecosystems 

in the world (Hicks 1997). Numerous wildlife species, 

including a diverse assemblage of birds, inhabit oakdominated forests. The songbird communities that 

depend on oak-dominated forests in the Southeast 

include a large number of neotropical migrants (Hamel 

et al. 1982; Thompson and Fritzell 1990). Since many 

of these species are declining, the management of their 

habitats has become a conservation priority. Although 

the prevailing songbird conservation paradigm in many 

eastern hardwood-dominated forests (especially large 

blocks of public forests) has been to limit harvests 

to single/group-tree selection or eliminate it entirely, 

thousands of acres of oak-dominated forests occur on 

private lands where wildlife conservation goals may be 

secondary to timber management priorities.

Game bird species also use oak stands in various stages 

of succession. The cultural and economic importance 

of maintaining harvestable populations of these 

species is a priority for many state agencies as well as 

private landowners. Prescribed fire in oak-shelterwood 

systems, when properly implemented, can provide 

the habitat diversity across the landscape that fosters 

healthy populations of game and nongame birds. The 

novel techniques suggested here provide an innovative 

management option that can satisfy the multiple goals 

of avian conservation and sustainable quality hardwood 

timber production.

Burning as a silvicultural technique in many forests 

traditionally has been associated with pine production. 

Conversely, it has been regarded as a force to keep 

out of hardwood forest management. The technique 

described here is an effective method for regenerating 

oak-dominated stands in the Southeastern Piedmont 

(Van Lear and Brose 1999). The steps involved in the 

process (partial harvest, burning, complete harvest) will 

create two-aged, open woodland, or shrubland habitats. 

Therefore, stands managed in different stages of the 

shelterwood-burn process across a landscape would offer 

habitats to forest-interior, edge-interior, open-woodland, 

and early-successional shrubland species. We believe that 

this technique has application beyond the Southeast 

to many other eastern forest uplands where oak is 

the desired dominant or codominant. The successful 

application of oak-shelterwood burn techniques may 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 175

be especially critical in areas such as the Southern 

Appalachian Mountains where a number of species 

require critical recovery or immediate management 

efforts. 

Caveats and Conclusions

Where landscape context and logistics allow, managing 

upland oak-hardwood stands using some of the options 

outlined here can potentially produce a variety of 

habitats that favor various game and nongame birds 

while effectively producing quality hardwood timber. 

Management and conservation priorities for birds will 

vary regionally. The silvicultural regime suggested here 

offers alternatives that can accommodate a variety of 

different regional and management contexts.

We have offered baseline information on the 

conservation scoring and management priorities for 

neotropical migrants and gamebirds using Partners in 

Flight rankings and our best inference as to the habitat 

occupancy of the silvicultural options described herein. 

While these scores provide a basis for understanding the 

relative importance of a particular management option 

for various species and suites of species, they should not 

be used as stand-alone prescriptions for management. 

Researchers and natural resource managers must take 

regional and site specific contexts into account. We 

suggest that the use of conservation value indexing 

(Twedt 2004; Nuttle et al 2003; Bryce et al. 2002), 

which incorporates abundance estimates and other 

demographic factors, may provide a more quantitatively 

rigorous assessment of habitat “value” that can be used to 

build robust scenarios for planning and management.

While wildlife and timber production goals frequently 

are in opposition, the ability to reliably reproduce oakdominated stands using a less intensive form of evenage management like the technique described here and 

associated options might be an effective tool for both 

wildlife conservation and sustainable timber production. 

As the demand for quality hardwood products increases, 

compromises regarding timber management and wildlife 

conservation in upland hardwood forests must be 

reached. While the oak-shelterwood burn system offers 

novel solutions for managing avifauna, the manager 

must always remember that he or she cannot manage 

every acre for every species. Rather, a knowledge of the 

landscape context and specific management objectives, 

supported by an adaptive management plan, offers 

opportunities for both the sustainable management of 

quality timber and diverse avian communities in eastern 

oak-dominated upland forests.

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1	 Fire 80 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

INFLUENCE OF FIRE ON MAMMALS IN EASTERN OAK FORESTS

Patrick D. Keyser1† and W. Mark Ford2

Abstract.—With the exception of small mammals, little research has been conducted in eastern oak 

forests on the influence of fire on mammals. Several studies have documented little or no change in 

relative abundance or community measures for non-volant small mammals in eastern oak (Quercus spp.) 

forests following fires despite reductions in leaf litter, small woody debris, and changes in understory and 

midstory composition and structure. Other studies have documented short-term improvements in quantity 

and quality of forage for white-tailed deer (Odocoileus virginianus), but none has demonstrated changes 

in population parameters. For other mammal species there are no explicit studies and influences must 

be based on inferences from other research. We suggest that species that prefer partially open canopies, 

herbaceous understories, reduced midstories, or savannah habitats likely will prosper in the presence of 

fire. Fire has the potential to both recruit as well as eliminate den sites and cavity trees; burning regimes 

and fire intensity likely will determine outcomes. Species of high conservation concern, such as the 

Indiana bat (Myotis sodalis) and some subspecies of fox squirrels (Sciurus niger), should benefit from 

increased fire in these landscapes. An insidious consequence of continued preclusion of fire in oak systems 

is the loss of structure and the change in vegetative species composition to more fire-intolerant species.

volant small mammals, typically relative abundance 

and species richness at the stand level or smaller, to 

fire in the Eastern United States (Masters et al. 1998; 

Ford et al. 1999; Keyser et al. 2001; Kilpatrick et al. 

2004). Incidental impacts of fire have been reported 

as well (Kirkland et al. 1996). Studies of economically 

important game species such as white-tailed deer 

(Odocoileus virginianus) have largely been confined 

to assessment of forage quality, availability, and use 

(Hallisey and Wood 1976; Wentworth 1986; Masters 

et al. 1993). Despite their high conservation status, fire 

impacts on nongame groups such as bats or charismatic 

mega-fauna such as black bears (Ursus americanus) 

have been examined only in conceptual, hypothetical 

approaches (Hamilton 1981; Carter et al. 2002). In this 

paper we review the literature relevant to fire effects on 

mammals in eastern oak forests. Our purpose is to draw 

meaningful inferences for species and species groups 

based on documented effects of prescribed fire or wildfire 

on habitat structure, and to discuss potential research 

directions to address current knowledge gaps. Fire or the 

lack thereof will continue to influence eastern oak forests 

and, in turn, the wildlife communities that depend 

on those forests for habitat (Kellison 1993; Brennan 

et al 1998). Accordingly, it is incumbent upon the 

natural resources community to work toward managing 

those outcomes to the benefit of sustainable wildlife 

populations.

INTRODUCTION

Fire has long been recognized as an important ecological 

driver and disturbance agent that shapes and maintains 

oak-dominated forest communities of eastern North 

America (Delcourt and Delcourt 1998; Frost 1998; Van 

Lear and Harlow 2002). Unlike other hardwood forest 

types, such as northern hardwoods and bottomland 

hardwoods, many oak-dominated types have relatively 

short fire return intervals (4 to 30 years) over most 

of the region (Frost 1998). A number of researchers 

have focused on concerns specifically related to the 

sustainability of oak forests (Clark 1993; Kellison 1993; 

Keyser et al. 1996; Brose et al. 1999; Brose et al. 2001), 

including those of the central hardwoods region (Van 

Lear et al. 2000; Sutherland and Hutchinson 2002).

Despite a number of studies on the effects of fire on 

wildlife in the Coastal Plain of the Southeastern United 

States, little work has been done on this subject in 

oak-dominated forests (Brennan et al. 1998; Lanham 

et al. 2002; Keyser et al. 2004), particularly as it 

relates to mammals (Keyser et al. 2001). Only several 

experiments have examined the response of non-

1

Forestry Division, MeadWestvaco Corporation, P.O. Box 577, 

Rupert, WV 25984; 2USDA Forest Service, Northern Research 

Station, Box 404, Parsons, WV 26287; †Corresponding author, 

304-392-1606, email: pdk2@meadwestvaco.com.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 181

NON-VOLANT SMALL MAMMALS

Despite the paucity of data on fire impacts on most 

mammals in eastern oak forests of North America, 

non-volant small mammals have been perhaps the best 

studied due to their abundance, diverse assemblages, and 

ease of capture. In coniferous forests of Western North 

America and along the mixedwood-boreal transition 

zone where natural stand-replacing fires and intense 

site-preparation burns are more common, impacts of 

fire on non-volant small mammals can be significant 

via direct mortality (Tevis 1956) and through habitat 

modifications that alter and shape species assemblages 

(Crete et al. 1995; Sullivan and Boateng 1996; Monroe 

et al. 2004). Nonetheless, Crete et al. (1995) noted that 

a mosaic of stand seres created by landscape-level fires 

added to overall wildlife diversity, including that of non-

volant small mammals.

In eastern oak forests, both natural and prescribed fires 

typically are of relative low intensity, are rarely stand 

replacing (Komarek 1974), and display site variability 

(Ford et al. 1999) at scales matching inherent variability 

of small mammal populations (Bowman et al. 2000). 

Consequently, negative impacts are not suspected in 

most cases. Orrock et al. (2000) noted that in some 

Appalachian habitats, small mammal species such as 

the southern red-backed vole (Clethrionomys gapperi) 

are specialized for mesic forest conditions and could be 

more fragmented and isolated by disturbance events. 

Results from both prescribed burning in the southern 

Appalachians of North Carolina (Ford et al. 1999) and 

a wildfire in the central Appalachians of Pennsylvania 

(Kirkland et al. 1996) suggest that small mammal 

abundance and species assemblages are not changed 

substantially by fire. Ford et al. (1999) noted that the 

abundance and assemblage patterns that existed prior 

to spring-burning persisted following burning with 

mesic-forest adapted species such as southern red-backed 

voles, smokey shrews (Sorex fumeus), and masked shrews 

(S. cinereus) more common in riparian areas and areas 

with abundant coarse woody debris in both burned and 

control stands. Moreover, data from Keyser et al. (2001) 

in central Virginia documented that the season of burn 

had little effect on non-volant small mammals with 

shrews (Sorex longirostris, S. hoyi, and Blarina carolinensis) 

and white-footed mice (Peromyscus leucopus) numbers 

not significantly different among oak shelterwood stands 

burned in the winter, spring, or summer, and unburned 

controls.

It is plausible that non-volant small mammals could 

be affected positively or negatively by fire in number 

of ways other than direct mortality. For example, on 

the Cumberland Plateau of Kentucky, prescribed fire 

in mixed mesophytic forests reduced the mass of soil 

invertebrates by approximately 35 percent (Kalisz and 

Powell 2000), which, in theory, could reduce available 

prey for Soricids. Conversely, Ahlgren (1966) observed 

that deer mice (Peromyscus maniculatus) responded 

positively to prescribed burning in Minnesota in 

part because fire consumption of the leaf litter and 

duff greatly increased seed exposure and, therefore, 

availability. Similar to forest floor structural changes 

observed in more intensive western fires, heavy 

reductions in leaf litter and downed coarse woody 

debris could negatively impact many non-volant small 

mammals in eastern oak forests. However, Kirkland et 

al. (1996) and Ford et al. (1999) observed virtually no 

change in coarse woody debris abundance following 

the low- to medium-intensity fires they studied. 

Additionally, substantial reductions in coarse woody 

debris following more intense fires in the region typically 

will be offset by overstory mortality and contributions 

to the future coarse woody debris pool (Regelbruggae 

and Smith 1994). Generally, though, data applicable 

to eastern oak forests are consistent with the findings 

of Kirkland et al. (1996) and Ford et al (1999) that 

ground-level vegetation change after one growing 

season following fire was less important than topoedaphic factors of slope and aspect (Wentworth 1986, 

Hutchinson et al. 2005). Prescribed fire in eastern oak 

forests can be effective in reducing woody midstory stem 

density (Brose et al. 2001), creating structural conditions 

that do benefit both southern flying squirrels (Glaucomys 

volans; Bendel and Gates 1987) and fox squirrels (Sciurus 

niger; Edwards et al. 2003).

Although wholly unknown, behavioral changes in 

small mammals during or following fire might cause 

an increase in vulnerability to predation. Rowan et 

al. (2005) radio-tracked eastern chipmunks (Tamias 
1	 Fire 82 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

striatus) following early spring fires in West Virginia on 

both burned and control plots. Despite fire intensities 

sufficient to consume most leaf litter, interrupt or modify 

the initial emergent patterns of herbaceous flora, and 

expose acorn caches, no difference was noted in mean 

home range size and habitat use between animals in the 

burned or control plots. Of the individuals captured 

and tagged within burned stands, none were observed 

leaving the burned areas or using adjacent unburned 

forest habitat that was in many cases less than 150 m 

away (Rowan et al. 2005). Home range overlap among 

conspecifics did not differ between burned and control 

plots, further suggesting minimal impact to eastern 

chipmunks.

BATS

Carter et al. (2002) provideed the most complete review 

of the relationship between fire and bats in eastern 

oak forests. Depending on the fire severity and fire 

susceptibility of tree species present, fire in eastern oak 

forest can both produce as well as consume standing 

snags that could be used as day-roosts by several bats 

species in the region (Starker 1934; Wendell and Smith 

1986; Moorman et al. 1999a,b; Abrams 2000). The 

endangered Indiana bat (Myotis sodalis) and the northern 

bat (M. septentrionalis) predominately use snags as dayroosts from spring through early fall (Menzel et al. 2001; 

Owen et al. 2002). Whereas little brown bats do roost in 

snags (Fenton and Barclay 1980), that species’ use is far 

less due in part to the availability of human structures 

as well as the overall change in forest condition, i.e., 

fewer large cavity trees and standing snags than in 

pre-settlement forests (Whitaker and Hamilton 1998). 

Although bats could be subject to direct mortality from 

the combustion of an occupied day-roost, especially 

one containing non-volant young in a maternity roost, 

most bats are capable of escaping a snag in advance of its 

consumption by fire. For bats that day-roost in foliage of 

live trees, e.g., hoary bats (Lasiurus cinereus) and eastern 

red bats (L. borealis), direct impacts probably result from 

temporary disturbance and displacement due to smoke 

(Rodrigue et al. 2001). Nonetheless, as nonhibernating 

migratory species, eastern red bats have been observed 

day-roosting in leaf and pine needle litter in upland pinehardwood forests of South Carolina and Arkansas where 

they were disturbed as winter burning occurred (Saugey 

et al. 1998; Moorman et al. 1999a,b). It is possible that 

eastern red bats might be unable to flee advancing fire if 

in a state of torpor while ground-roosting.

Realistically, however, species that depend on snags 

probably benefit from a net addition of snags in 

burned stands in a region where it is believed that snag 

abundance is below that needed for many wildlife 

species (Carey 1983). For example, Wentworth (1986) 

noted that midstory tree mortality was pronounced on 

variable winter burns in the southern Appalachians, 

conditions that likely produce a net gain of day-roosts 

highly suitable for northern bats (Owen et al. 2002). 

Still, there are virtually no data on the net gain or loss or 

longevity and useful to bats of larger snags following fire 

in eastern oak forests. Lastly, Carter et al. (2002) noted 

that bats wintering in caves and mines might be subject 

to negative impacts from smoke in areas where air-flow 

into caves or mines occurs. However, few instances of 

this have been documented and none with recorded 

bat mortalities (D. Krusac, USDA Forest Service, pers. 

commun.).

Relative to all other mammalian taxa in the eastern 

oak forest, knowledge gaps in Indiana bat habitat 

management in conjunction with fire probably are 

the most critical to address. Maternity roost habitat 

of Indiana bats often is characterized by stands with 

numerous snags and relatively low overstory canopy 

cover (Kurta 2004), indicative of conditions following 

flood events in bottomland forest habitats in the 

Midwest (Carter and Feldhamer 2005) or following 

moderate to severe fires in upland habitats in the 

Appalachians (Beverly and Gumbert 2004). Although 

less demanding of particular thermal conditions (i.e., 

warm temperatures and direct solar radiation) helpful 

for the growth and development of juvenile bats, male 

Indiana bats also readily use day-roosts in snags created 

following prescribed burning on the Cumberland 

Plateau of Kentucky (J. MacGregor, Kentucky Wildlife 

and Fisheries Resources Commission, pers. commun.). 

Similarly, we have documented two male Indiana bats 

day-roosting in burned forest habitats in the Allegheny 

Mountains of West Virginia where diameter-limit 

harvests followed by early spring wildfire produced open 

conditions (20 to 50 percent canopy cover) and copious 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 183

small to medium snags (< 40 cm d.b.h.) with large 

amounts of exfoliating bark. At that same site, Beverly 

and Gumbert (2004) described an Indiana bat maternity 

roost occurring in a small fire-killed red maple (Acer 

rubrum) snag (31 cm d.b.h.). This roost occurred at an 

elevation of 917 m where conditions were considered 

too cool during the growing season to support maternity 

activity (Brack et al. 2002). Because that roost was near 

a known winter hibernacula (< 1,000 m) and several 

male roosts in fire-killed sugar maple (A. saccharum ) 

and red maple (unpublished), it has been hypothesized 

that Indiana bats in the Appalachians probably formed 

transitory maternity roosts within a shifting mosaic of 

habitats that may have followed disturbances such as 

fire across the landscape (T. Carter, Southern Illinois 

University, pers. commun.).

Fire effects on the structure of bat foraging habitat and 

fire impacts to bat prey resources are largely unknown 

(Carter et al. 2002). Menzel et al. (2005) and Ford et 

al. (2005) both hypothesized that forest management 

activities that decreased stem density and or created 

canopy gaps and openings would benefit several bat 

species in the east that are adapted to foraging in less 

cluttered conditions, e.g., big brown bat (Eptesicus fuscus) 

and hoary bat. If they are correct, prescribed fire could 

be a useful tool for enhancing foraging habitat for both 

species by creating a more complex canopy structure 

through mortality of individual canopy stems.

WHITE-TAILED DEER

Stransky and Harlow (1981) provided a thorough review 

of the literature regarding deer habitat and fire. Because 

there were few studies at that time directly examining 

deer and fire relationships, theirs was a thorough review 

of work on forage, including that for livestock, and fire 

effects on vegetation in order to draw inferences relating 

to the likely effect of fire on deer habitat. Stransky and 

Harlow concluded that winter burns increased forage 

crude protein and phosphorous (P) content; increased 

the number of woody plant stems; increased the coverage 

of grasses, forbs, and legumes; and reduced soft mast 

production. Most of these effects were short term (1 to 

3 years). Infrequent growing season burns had similar 

effects except that woody stems were reduced. However, 

frequent summer burns would eventually eliminate 

woody stems and lead to domination of the site by 

grasses and fire-adapted forbs. These structural changes 

were established empirically for pine forest systems 

by the long-term (43 years) experiment conducted by 

Waldrop et al. (1992).

Wood (1988) and Wentworth (1986) each documented 

limited responses of vegetation to burning, albeit 

generally consistent with the foregoing summary 

of Stransky and Harlow (1981). Both studies were 

conducted in closed-canopy mature forests. Hurst et al. 

(1980) and Masters et al. (1993) conducted experiments 

in which they documented three- and sevenfold increases 

in forage production. The opening of canopies in both 

studies was critical to these responses with fire modifying 

plant communities released by canopy disturbance. 

Working in a scrub oak community in the Appalachians, 

Hallisey and Wood (1976) found that crude protein, P, 

and amount of forage generally were increased by fire. 

In another experiment, Brennan et al. (1998) precluded 

fire from a previously burned stand and reported a 

dramatic decrease in herbaceous cover largely due to 

encroachment by fire-intolerant woody vegetation. 

With the exception of Wentworth’s (1986) and Hallisey 

and Wood’s (1976) studies, all of the work cited was 

conducted in pine or pine-hardwood systems.

Two studies (Hobbs and Spowart 1984 and Main 

and Richardson 2002), neither conducted in eastern 

hardwoods, empirically documented behavioral changes 

(habitat use and foraging, respectively) of ungulates in 

response to fire. Main and Richardson (2002) deployed 

infrared triggered cameras in areas that had been burned 

6, 24, and 48 months previously to document use by 

white-tailed deer. They reported differences between the 

48- and 24-month postburn sites as well as between the 

48- and 6-month postburn sites, with use by deer 3 to 

4 times greater in the more recently burned sites. They 

concluded that deer were attracted to the availability of 

better forage in the more recently burned areas. Hobbs 

and Spowart (1984) reported only modest improvements 

in site-level forage quality, but substantial improvements 

to actual diets of both mountain sheep (Ovis canadensis) 

and mule deer (Odocoileus hemionus) on burned versus 

unburned sites in Colorado. They determined that 

due to selective foraging, these ruminants were able to 
1	 Fire 84 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

increase both CP and in vitro digestible organic matter 

by 100 to 400 percent. They contend that studies that 

look only at plant responses underestimate the impact of 

burns on overall diet quality.

From the foregoing discussion it seems clear that where 

some canopy disturbance is present, fire alters forage 

quality and quantity, deer behavior, and likely deer 

diets. Unfortunately, no study has linked any population 

parameter to fire in a conclusive manner. Johnson et al. 

(1992) documented subtle, short-term improvements 

in white-tailed deer body mass and condition after a 

single large (18,200 ha) wildfire in the Coastal Plain 

of North Carolina. However, cause-and-effect data 

for fecundity, growth, and survival for deer on burned 

habitats are lacking. Given the degree to which whitetailed deer herds exceed cultural carrying capacity 

(society’s tolerance threshold for a species) despite being 

below biological carrying capacity throughout much of 

this region (Keyser et al., in press), would suggest that 

ample nutrition is available to support large and healthy 

populations in the absence of fire-enhanced habitats.

An insidious impact of fire suppression on deer habitat 

in the region is the possible loss of oak mast through 

a marked reduction of the oak component in these 

forests. Mast is an important and preferred food source 

for deer (Harlow and Hooper 1972; Osborne et al. 

1992; Wentworth et al. 1992), one that is widespread 

and often available in association with excellent cover. 

It is difficult to predict the impact of a large-scale loss 

of this resource on regional deer populations, but it is 

potentially profound. This would be especially true in 

landscapes that are heavily forested and lack agronomic 

forages that could easily offset the loss of high energy 

and highly palatable forage provided by acorns.

OTHER SPECIES

As is the case with deer, small mammals, and bat 

communities, little work has been done on fire effects 

on other mammal species that are associated with 

eastern oak forests. Both Hamilton (1981) and Weaver 

(2000) discussed fire influences on black bear but were 

confined to inferring effects on habitat because of a 

lack of published data. They concluded that periodic 

use of fire could be positive in terms of enhancing 

soft mast production and maintaining herbaceous 

plant communities. Fire can be important in both the 

formation and destruction of den trees, though we are 

unaware of any work that documents optimum regimes 

for recruitment and/or maintenance of cavity trees in 

such areas. In the Appalachians, bears frequently rely on 

accumulations of large woody debris for ground dens 

(Quince 2002), which also can be eliminated by fire. 

Thus, fire can have both positive and negative effects on 

availability of dens. Where fires provide broken canopies 

and more diverse midstory and understory development, 

particularly as it relates to soft mast and herbaceous 

vegetation, fire likely would result in positive changes 

in habitat quality. On the other hand, to the extent that 

fire negatively affects mast production, it may negatively 

effect habitat quality. We do not believe that fires intense 

or frequent enough to induce such changes are likely to 

be the norm in most eastern oak systems.

Much of the foregoing discussion about black bear 

denning habitat likely is applicable to raccoons (Procyon 

lotor). Owen (2003) documented use of both ground 

dens and tree cavities by raccoons in the central 

Appalachians, though maternity dens were almost 

exclusively in tree cavities. Thus, burning frequencies 

and intensities that maintain adequate snags appear 

to be important for this species. Although conducted 

in the Coastal Plain, Jones et al. (2004) documented 

a reduction in use of forests burned the previous year 

versus unburned stands by raccoons. It is difficult to 

infer how this might apply to upland hardwood systems. 

As with black bears, fire effects that enhance soft or 

hard mast production likely would increase raccoon use 

(Owen 2003).

We are aware of no literature that deals specifically with 

responses of gray (Sciurus carolinensis) or fox squirrels 

to fire, a circumstance that has not changed since 

Kirkpatrick and Mosby (1981) reviewed the literature. 

Although we are in agreement with them that fire is 

unlikely to positively affect gray squirrel habitat, it may 

have important implications for conservation of rare 

or threatened subspecies of fox squirrels. Fox squirrels 

prefer more open stand structure and greater understory 

development (Edwards et al. 2003), conditions that 

could be created through prescribed burning.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 185

Elk (Cervus elaphus) may benefit from the increased 

use of fire in eastern forests due to their preference for 

herbaceous understories. Masters et al. (1997) reported 

increased use of burned areas by elk in a pine hardwood 

system in the Ouachita Mountains where herbaceous 

cover had been developed by repeated fires. Elk rely on 

a wide variety of forages, but grasses and forbs dominate 

elk diets on a year-round basis and in most habitats 

(Boyd 1978). Most elk restoration efforts in the Eastern 

United States focus on release sites with adequate 

herbaceous habitats (McClafferty and Parkhurst 2001). 

As interest in restoration of elk increases in eastern 

states and as elk populations expand in Pennsylvania, 

Kentucky, North Carolina, and Tennessee, managers 

should consider the role of fire in developing oak 

savannah habitats.

SUMMARY AND CONCLUSIONS

The state of our current understanding of fire effects on 

mammals in the oak forests of Eastern North America 

remains largely based on inference from work conducted 

in other regions or on indirect effects of fire on various 

habitat components. On the basis of these studies, it is 

apparent that fire can have a beneficial impact on habitat 

quality for a number of species of mammals. Were fire 

to be applied on a broad enough scale, these habitat 

enhancements could reasonably be expected to have a 

positive effect on population parameters as well. It is 

unclear in today’s social and political climate that land 

managers will apply fire at such a scale. Conversely, it 

could be expected that on a localized scale fire could have 

a significant impact on local populations of mammals, 

either behaviorally or numerically. Thus, individual 

landowners or managers interested in enhancing habitat 

for deer could use prescribed fire as a tool.

However, the continued exclusion of fire from most 

landscapes in eastern oak forests may have serious 

implications for the continued presence of oaks as a 

major component of regional forests. It is this scenario 

that likely will have broader impacts on mammalian 

communities of the region going forward. Current 

trends in forest communities suggest that oaks will be 

a diminishing part of our future forests. Such a change 

could have a deleterious effect on numerous species of 

wildlife.

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Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 191

Obstacles and Solutions 

to Using Prescribed Fire 

in Oak Forests
1	 Fire 92 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Prescribed fire: What influences public approval?

Sarah M. McCaffrey1

Abstract.—Except in remote areas, most prescribed fires will have some effect on members of the public. It 

is therefore important for land managers to work with the public before, during, and after a prescribed burn. 

To do this effectively, managers need to have an accurate idea of what people do and do not think about 

prescribed fire and they need to understand what shapes those opinions. This paper summarizes findings 

from recent research studies on the social acceptability of prescribed burns and identifies the key factors that 

people consider in forming their opinions of prescribed fire. Results indicate that there is a fairly high level 

of public acceptance for use of prescribed fire and that smoke, concerns about escape, and trust are key issues 

shaping that support. In addition, there is a clear link between understanding of the purpose and intended 

benefits of prescribed fire and approval of its use. The lesson for managers who wish to introduce prescribed 

fire in their communities is that they are most likely to gain public support if they: 1) increase familiarity 

with the practice; and 2) work to build trust between officials from the implementing agency and the public 

information about these studies is summarized in Table 

1. Some of the studies have been completed while 

others are still in progress; information about them is 

drawn from published articles, project reports, and, in 

one case (See Table 1, McCaffrey 2005), directly from 

focus group transcripts. Although there is local variation 

in forest composition in the study areas3,the findings 

are reasonably consistent across diverse ecosystems and 

different regions of the country. This paper should 

therefore provide managers with a sense of the basic 

dynamics that shape public opinions of prescribed fire 

to help guide development of programs that fit local 

circumstances.

Approval

Prescribed burning is a largely acceptable practice with 

roughly 80-90% percent of respondents across studies 

finding it an appropriate management tool (Bright and 

Carroll 2004; Cortner et al. 1984,;McCaffrey 2002; 

Shelby and Speaker 1990; Shindler et al. 1996). In those 

surveys that explored strength of support, roughly 30 

percent of respondents gave strong approval for use of 

Introduction

I think what happens when we have prescribed 

burns is the majority of the people say “Well, that’s 

something that has to be done.” And there’s a 

minority of the people that complain about it, but 

they get their names in the paper.2

(Hamilton focus group participant)

Prescribed burning is a key tool for managers working to 

reduce fuel loads or restore fire adapted ecosystems. Yet 

it can also be a problematic practice that the public may 

not accept or support. Since prescribed fires in all but 

the most remote areas will have some effect on members 

of the public, it is important for managers to work with 

the public before, during, and after a prescribed burn. To 

do this well, it is useful to have an accurate idea of what 

people do and do not think about prescribed fire and 

what shapes those perceptions. As the introductory quote 

suggests, public views of prescribed fire are generally 

more sophisticated and less negative than managers 

might expect.

This paper will discuss findings from recent studies 

(most sponsored by the National Fire Plan) about the 

social acceptability of prescribed fire, the key variables 

that influence approval or disapproval, and the roles 

that those variables play in shaping opinion. Basic 

1

USDA Forest Service, Northern Research Station, 1033 

University Place, Suite 360, Evanston, IL 60201-3172, 

847-866-9311 ext. 20, email: smccaffrey@fs.fed.us.

2

All quotes are taken from participants in a series of focus 

groups held to examine public views of various aspects of fire 

management (McCaffrey 2005, Table 1,). A total of fifteen 

focus groups were held in five different fire prone locations 

in the Western United States. Focus groups were made up 

of randomly selected local members of the public and were 

transcribed verbatim.

3

Only the California and Missouri sites of the Winter et al. 

2005 study included some oak woodlands.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 193

prescribed burning and another 50 percent gave qualified 

approval (Blanchard 2003; Brunson and Evans 2005, 

Shindler and Toman 2003, Shindler et al. 2003; Winter 

et al. 2005).

Smoke

Smoke is often considered a major barrier to use of 

prescribed fire. This is a reasonable expectation given 

that smoke is a health problem for roughly 30 percent 

of study households (Blanchard 2003; McCaffrey 2002; 

Shindler et al. 1996; Winter et al. 2005). However, 

in their four state study, Winter et al. (2005) found 

that smoke was significantly related to prescribed fire 

attitudes in only one site, Missouri, where the belief that 

prescribed burning meant more smoke now and less later 

was positively related to approval. In general, people 

appear to understand that no smoke is an unrealistic 

option: they will be exposed to smoke, either from a 

wildfire or from a prescribed burn, and so long-term 

trade-offs will need to be made. One way to manage 

the health issues is to provide adequate warning of a 

prescribed burn allows those with health issues to make 

arrangements.

I think they would (tolerate smoke), if it is 

communicated ahead of time so that asthmatics could 

stay inside; like we do now when they broadcast 

something that there’s a wildfire. You can plan ahead. 

(Reno focus group participant)

Maybe if it had advance notice, they would know for 

how long it would take and what the purpose was. 

(Boulder focus group participant)

Focus groups in Washington found that, although people 

often don’t differentiate, the source of the smoke can 

influence approval (Weisshaupt et al. 2005). Members of 

an anti-smoke group remained opposed to smoke from 

agricultural burning because all benefits went to the 

farmer but were more open to smoke from prescribed 

fire because the benefits of burning accrued to all. Many 

participants, including those in the anti-smoke group, 

were willing to make trade-offs between some “managed” 

smoke now in return for less smoke from future 

wildfires.

Topography is a local variable that can influence 

acceptability of prescribed fire due to smoke concerns. 

Study Where Who Method 

Blanchard & 

Ryan 2002

Within a two mile radius of 

Myles Standish State Forest in 

Massachusetts

Seasonal and year round residents Mail survey

Bright and 

Carroll 2004

Front Range, Colorado, Southern 

Illinois, Chicago Metropolitan area

Residents near National Forests and 

random Chicago households Mail survey

Brunson and 

Evans 2005

Wasatch, Utah, and Salt Lake 

counties

Residents (including 113 who had 

been answered the same survey two 

years earlier)

Longitudinal 

mail survey

McCaffrey 2005

Flagstaff, AZ; Boulder, CO; 

Hamilton, MT; Reno, NV; San 

Bernardino, CA General members of the public Focus groups

Shindler et al. 

2003

Forest communities adjacent to 

National Forests in Wisconsin, 

Michigan, and Minnesota

Residents Mail survey

Weisshaupt et al. 

2005 Missoula, MT; Spokane, WA Native Americans, urban and rural 

residents and an anti-smoke group Focus groups

Winter et al. 

2005, 2004, 2002

California, Florida, Michigan and 

Missouri Homeowners near forested lands Focus groups 

and mail survey

Table 1.
1	 Fire 94 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

People who live in areas prone to inversions or in 

valleys that “collect” smoke from other areas are more 

sensitive to smoke issues and may find prescribed fire 

less acceptable. This dynamic is demonstrated by the 

following excerpt from a Reno focus group.

Ann: I don’t think very many people 

would tolerate it. My Grandma, she 

has problems like that. When there’s 

smoke, she can’t tolerate it. A lot of 

people can’t tolerate it.

Barbara: And we are in a valley, so it all just 

sits here.

Carl Inversion.

Carol: Even when they do a controlled 

prescribed burn in Yosemite, it still 

comes here.

But individuals in these areas also may recognize that 

topography means they have little control over the smoke 

from wildfires but some control over that of prescribed 

fire, as in Hamilton.

Come August the whole valley is going to be filled 

with smoke anyway. If they can do something 

productive and burn away some of that slash that 

might slow down the fire a bit, then I’m all for it.

Control

Concern over a burn getting out of control is another 

major issue. Early studies found that fear of a prescribed 

burn escaping influenced approval (Cortner 1984; 

Shelby and Speaker 1990). Seventy percent of the 

respondents in Shindler et al.’s study (2003) indicated 

they were moderately to greatly concerned about such 

a possibility. Winter et al. (2005) found that the belief 

that prescribed fire would lead to uncontrolled fires was 

negatively related to acceptance, the only outcome belief 

significant across all four study sites.

In Utah, Brunson and Evans (2005) compared responses 

from individuals surveyed in 2001 and again in 2003 

after an escaped burn (Cascade II) occurred in the area. 

They found that, after the escape, respondents expressed 

significantly increased concern (from 19 to 44 percent) 

about a prescribed fire taking place within 10 miles of 

their home suggesting that the escape may have led some 

people to have lower confidence about the ability to keep 

a prescribed fire under control. Notably, although almost 

half of the respondents stated that they held a more 

negative view of prescribed fire after the escape, in reality, 

when compared with responses from before the escape, 

their judgements of the acceptability of prescribed fire 

had not changed significantly. What had changed was 

an increased concern about the health impacts of smoke 

and decreased belief that smoke could be managed 

acceptably. Even with these changes only 13 percent 

thought smoke made prescribed fire not worth using.

Part of the issue with concerns about escaped burns 

is that they are what grabs people’s attention, not the 

successful burns—this creates a rather small sample 

upon which people base their opinions. In the series of 

focus groups throughout the West (McCaffrey 2005, 

Table 1), conversations about prescribed fire usually 

started with an instant negative reaction related to 

escapes but, in several cases, discussion then evolved 

around the question of how many problem burns there 

actually were in a given year compared to non problem 

burns. When people concluded that escapes were a very 

small percentage of total burns they were much more 

comfortable with them. They suggested that more effort 

needs to be made to publicize all prescribed burns to 

provide some perspective on the relative number of 

escaped burns. 

I think we need to know more. Just like John said, 

if 90% of them are successful, we need to know 

about it. But we just hear about the ones that aren’t. 

(Reno focus group participant)

The only time you hear it is always the bad fires. I 

think that they (prescribed burns) are helping us a 

lot. I would say approximately 5% goes out of hand. 

(San Bernardino focus group participant)

Trust

Trust in the agency administering the burn is the 

remaining key variable that shapes public acceptance 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 195

of prescribed fire. As one Reno focus group member 

said when asked whether people would tolerate more 

prescribed burning: “more if you trust the guy that starts 

it.” Winter et al. (2005) found that trust in government 

was a significant predictor of intention to approve 

prescribed burning in all four study sites. In Missouri, 

which had the highest trust levels of the four states, trust 

also had the largest effect on attitudes. Perhaps the most 

important impact of the escaped burn in Utah was not 

its effect on public views of prescribed burning but on 

trust levels which decreased significantly for both the 

U.S. Forest Service and the Bureau of Land Management 

but not for state, county, or local government agencies, 

with the local rural residents showing lower trust levels 

than the metropolitan sample (Brunson and Evans 

2005). For three of their study sites (California, Florida, 

and Michigan), Winter et al. (2004) examined what 

elements were associated with trust and found that the 

strongest association for all three sites involved views on 

agency competence.

Other variables

A variety of other issues such as past experience, wildlife 

concerns, and aesthetics are thought by many managers 

to influence approval of prescribed fire. However, results 

from the referenced studies do not show as strong or as 

consistent an effect as smoke, control, and trust. Most 

studies found no significant relationships with these 

three variables and those that did did not necessarily find 

a dependable effect. For instance, in terms of experience, 

Blanchard and Ryan (2004) found that individuals with 

past personal experience of wildland fire had a higher 

level of support for use of prescribed fire than those who 

had not. Conversely, Winter et al. (2005) found that past 

experience with wildland fire or prescribed fire was not a 

significant explainer of attitude.

Studies that explored how concerns about wildlife 

shaped acceptability also showed mixed results. Almost 

half of Shindler et al.’s (2003) respondents expressed 

at least moderate concern about loss of wildlife and 

fish habitat from a prescribed burn but 68 percent 

also thought it improved wildlife habitat. Winter et al. 

(2005) found that belief that prescribed fire improved 

wildlife conditions was positively related with approval 

in two of their sites: California and Michigan. In 

terms of aesthetics, while 42 percent of respondents 

in Minnesota, Wisconsin, and Michigan expressed at 

least moderate concern about scenic quality, only 14 

percent felt prescribed fire impacts on scenic quality 

were unacceptable (Shindler et al. 2003). Winter et al. 

(2005) found a weak association between aesthetics and 

approval for the California site and no association for the 

other three sites.

Finally, one item to consider that was examined in only 

one of the studies (Winter et al. 2005) is the belief that 

use of prescribed fire reduces firefighting costs in the 

long run—which was positively related with attitudes in 

California, Michigan, and Florida.

Understanding

The most consistently found relationship in the studies 

is the concept that familiarity with a practice leads to 

acceptance. That knowledge and familiarity with a 

practice is associated with increased support for fuels 

management practices fits with findings from earlier 

wildfire studies (Carpenter et al. 1986; Gardner and 

Cortner 1988; Loomis 2001; McCaffrey 2002). More 

recent studies have also found a strong link between 

knowledge and support for a treatment method, whether 

prescribed fire or thinning. Shindler et al. (2003) 

found that support for both treatment methods was 

significantly associated with the respondent’s natural 

resource knowledge—the greater the knowledge the 

greater the support as well as the greater the confidence 

in the U.S. Forest Service. Of the three states surveyed, 

Minnesotans were the best informed and the most 

tolerant of fuel treatments while Michiganders were 

least informed and least supportive. In Massachusetts, 

Blanchard and Ryan (2004) found knowledge levels 

to be the most significant factor determining support 

for prescribed fire; with a higher level of knowledge of 

prescribed burning significantly associated with increased 

support for its use as well as lower concern with related 

risks. Those with some knowledge of prescribed burning 

were less likely to think it was too dangerous a practice to 

be used, to be concerned about prescribed burning near 

a home, and to be concerned about smoke, appearance, 

and the effects on animals and their habitat.
1	 Fire 96 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

In Florida, the state in the Winter et al. study (2005) 

where prescribed fires were most common, respondents 

had the highest approval rate for prescribed burning 

and, as previously discussed, also had the highest level of 

trust in government agencies doing prescribed burning. 

They also were more likely to think that prescribed 

burns restored more natural conditions and improved 

conditions for wildlife. In the Washington focus groups, 

tolerance for prescribed burning increased, particularly 

amongst members of the anti-smoke group, as 

participants learned new information about the practice 

during discussion (Weisshaupt et al. 2005).

Understanding the ecological benefits of prescribed 

burning appears to be particularly important in shaping 

approval. Carpenter et al. (1986) reviewed three previous 

studies and found that acknowledgement of beneficial 

effects was the most “pervasive” influence in approving 

various fire management methods. More recently, 

Winter et al. (2005) found that belief that prescribed 

burning restored wildlands was positively associated with 

attitudes toward prescribed fire in Missouri as well as in 

Michigan (albeit at a weaker level), while, as indicated 

earlier, beliefs that prescribed fire improved wildlife 

conditions was positively associated with attitude toward 

the practice in California and Michigan.

Understanding ecological benefits can also make smoke 

less of a concern. In the Washington State focus groups, 

participants became more tolerant of smoke from a burn 

as they understood the beneficial effects of prescribed 

fire (Weisshaupt et al.2005). Shindler found that 2/3 of 

respondents in Oregon agreed that smoke was acceptable 

if it helped forest health (Shindler et al. 1996). As one 

Hamilton focus group participant responded when asked 

if people would tolerate smoke from increased use of 

prescribed burns: “I think most people would tolerate it, 

if they think, they knew it was good.”

Caveats

Results from the referenced studies thus indicate that, 

at a general level, there is a fairly high level of public 

acceptance for use of prescribed fire and that smoke, 

control and trust are key issues shaping that support. 

However, several caveats are important to keep in mind 

in applying this knowledge.

Local context matters

Local context, such as history and cultural practices, 

needs to be taken into account as it can have a significant 

effect on specific attitudes. For instance, Winter et al. 

(2002) found two exceptions to the general pattern 

of around 30 percent strong approval for prescribed 

burning: in Florida 40 percent were extremely positive 

about prescribed burning whereas in Michigan a mere 

10 % were extremely positive. This last is generally 

attributed to a 1980 prescribed fire that escaped and 

killed a firefighter, destroyed 44 houses, and is still 

discussed. Trust in government was also much lower in 

Michigan where only 27 percent of respondents trusted 

the government to make proper decisions about use of 

prescribed burning, as compared to 55 percent in Florida 

and 46 percent in California.

Avoid Preconceived Notions

It also is important to be careful of preconceived 

notions. Bright and Carroll’s (2004) study found very 

few significant differences between three groups often 

thought to hold different views: homeowners near 

National Forests along the Front Range of Colorado 

(classic wildland-urban interface or WUI) and Southern 

Illinois (very rural) and residents of the Chicago 

Metropolitan region. For all three groups, the most 

important factor in positive support for prescribed fire 

was if there was a recent history of fire and the second 

most important factor was if the burn was in a remote 

areas. The only major difference between the locations 

was in how primary use of the forest shaped acceptability. 

Illinois residents from both locations found prescribed 

fire more acceptable if the primary use of the land was 

commercial, while Front Range residents found it more 

acceptable if the primary use was recreational. However, 

primary use was less important in shaping acceptability 

than current conditions and forest location. The fact 

that such disparate areas have largely similar views is one 

example of how dichotomies, such as urban-rural and 

WUI versus non-WUI, can be misleading in terms of 

understanding public beliefs on fire management.

There also may be expectations that various sociodemographic characteristics, such as income or age, will 

be associated with specific attitudes. Although some 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 197

studies have found socio-demographic variables that were 

tied to attitudes, there has been no clear or consistent 

pattern.

Understanding is a two way street

Finally, the fact that there is a clear link between 

familiarity with a practice and acceptance does not 

mean that increasing acceptance of prescribed fire is 

simply a case of providing information. Shindler’s survey 

of Wisconsin, Minnesota, and Michigan showed that 

the most trustworthy and most helpful methods of 

information dissemination were guided field trips and 

interaction with agency personnel. Such interactive 

methods are most effective at changing attitudes and 

behavior as they allow people to question and clarify new 

information (Monroe et al. 2005). Manager’s in turn can 

learn through this process about key public concerns and 

issues and tailor their management efforts to account for 

them.

Conclusion

Contrary to the expectations of many managers, 

prescribed fire is a largely acceptable practice where 

objections to sensitive issues such as smoke and loss 

of control can potentially be overcome with dialogue 

as understanding of purpose and benefits increases 

tolerance. Thus managers introducing prescribed fire 

as a new tool may not have immediate acceptance but 

likely can look forward to increased public acceptance 

and support as people become more familiar with 

the practice. However, this increased acceptance is 

not automatic. Trust in agency implementation is 

also important. Even with good knowledge, low trust 

levels will likely mean low tolerance for prescribed fire. 

Fortunately, the very dialogue that agencies engage in to 

build knowledge bases can also help build relationships 

and trust.

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PA: U.S. Department of Agriculture, Forest Service, 

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of homeowner’s perceptions of wildland fire 

hazards: A case study from southern California.

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Prescribed fire and public support: knowledge 

gained, attitudes changed in Florida. Journal of 

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conceptions of wildfire risk. In: Martin, W.; Raish, 
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C.; Kent, B., eds. Wildfire risk: human perceptions 

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a synthesis of research related to communicating 

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Station. 42 p.

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David V., eds. Natural and prescribed fire in Pacific 

northwest forests. Corvallis, OR: Oregon State 

University Press: 253-259.

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acceptance of forest conditions and fuel reduction 

practices: a survey of citizens in communities 

adjacent to national forests in Minnesota, 

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Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 199

Top Ten Smoke Management Questions for Fire in Eastern

Oak Forests

Joseph J. Charney1†, Ann L. Acheson2, and Andrea Stacy3

visibility is reduced and the likelihood of traffic accidents 

increases. Ozone, which is formed through atmospheric 

reactions of nitrogen oxides and volatile organic 

compounds, is also a respiratory irritant.

As population centers surrounding eastern oak forests 

expand and encroach on these ecosystems, increasing 

the wildland/urban interface, smoke management issues 

will become increasingly important and increasingly 

complex. In some situations, smoke management could 

become the determining factor in “go/no go” decisions 

for prescribed burning.

The purpose of this paper is to outline pertinent smoke 

management questions that fire and smoke managers 

in the Central Hardwoods will face in the near future. 

The next section identifies and describes the “Top 

Ten” management questions that burn planners should 

address when implementing burning programs in 

the region. Section 3 will then describe the smoke 

management tools that are currently available, or that 

will be available in the near future, to help fire and 

smoke managers address these questions when preparing 

for prescribed burns. Section 4 includes a discussion and 

some concluding points.

TOP TEN SMOKE MANAGEMENT 

QUESTIONS

This section presents what we feel are the Top Ten smoke 

management questions that will impact prescribed burn 

decisions in eastern oak forests in the near future.

10. Where are eastern oak forests—Where will 

burning occur?

The Central Hardwoods (Fig. 1) cover an area from 

northern Georgia, west to Arkansas, northeast to the 

Great Lakes and New England, and then south along 

the eastern slopes of the Appalachians (Stringer and 

Loftis 1999). This area is predominantly oak-dominated 

deciduous forests in hilly-to-mountainous areas. The 

INTRODUCTION

In a time when fire planners and forest supervisors in 

many parts of the United States are implementing plans 

to increase the number and frequency of prescribed 

burns, the issue of smoke impacts on air quality 

is becoming increasingly important. Smoke from 

prescribed fires can have a dramatic impact on air quality 

and visibility on local, regional, and national scales, with 

corresponding temporal scales ranging from hours to 

days to months. As new national air quality regulations 

are implemented, smoke management will become 

an increasingly important concern when planning 

prescribed burns in the Eastern Oak Forests.

In the Eastern Oak Forests, prescribed fire is becoming a 

more common practice in ecosystem management (Ward 

and Brose 2004). Although there are many resource 

benefits associated with prescribed fire, the potential for 

the resultant smoke to impact human health and public 

safety also needs to be addressed. Wildland fires release 

various pollutants, including, but not limited to, fine 

particulate matter, nitrogen oxides (which are precursors 

to ozone formation and, thus, a surrogate for ozone), 

volatile organic compounds (also precursors to ozone 

formation), and carbon monoxide. However, in terms of 

prescribed fire, the pollutants of major concern are the 

fine particulates, referred to as PM2.5. About 70 percent 

of particulates released by wildland fires are within the 

fine particle size class (particles less than 2.5 microns 

in diameter) (Hardy et al. 2002). Elevated levels of fine 

particulates are dangerous because they can penetrate 

deep into human lungs and increase the risk of serious 

health problems, especially for those with respiratory 

illnesses. Furthermore, when smoke plumes with high 

particulate concentrations intersect roads and highways, 

1

USDA Forest Service, Northern Research Station, Stephen 

S. Nisbet Bldg., 1407 S. Harrison Road, Room 220, East 

Lansing, MI 48823; 2USDA Forest Service, Region 9, Fire and 

Aviation Management, 27750 State Route 7, Marietta, OH 

45750; 3USDA Forest Service, Monongahela National Forest, 

200 Sycamore St., Elkins, WV 26241; †

Corresponding author, 

517-355-7740 ext. 105, email: jcharney@fs.fed.us.
2	 Fire 00 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Central Hardwoods cover approximately 125 million 

acres in eleven states, making it the most extensive 

temperate deciduous forest in the world.

9. Where are the people—what percentage of people in 

the US live within the vicinity of eastern oak forests?

Approximately 65 million people live in the states east 

of the Mississippi (U.S. Bureau of the Census 2000), 

with the highest population density along the Eastern 

Seaboard, the Great Lakes coast, and the southern tip of 

the Appalachians (Fig. 2). Approximately 25 percent of 

the population of the United States lives in the Central 

Hardwoods states, and over 90 percent of the Central 

Hardwood Forest is owned by private interests, primarily 

non-industrial forest owners (Hicks 1997).

8. Where are the roads?

One of the major concerns with smoke management is 

whether the smoke from a prescribed burn will impact 

a roadway. The highest density of interstate highways 

and primary and secondary roads in the Unites States 

occurs east of the Mississippi (Fig. 3). A major source of 

complexity in eastern smoke management and prescribed 

burning is the fact that a road of some sort almost 

invariably occurs within the immediate locale of a burn 

site.

7. Where are existing air pollution problem areas?

Outside of California, the eastern United States has 

the greatest number of areas not meeting the National 

Ambient Air Quality Standards (NAAQS) for a variety 

of criteria pollutants (U.S. EPA 2006): sulfur dioxide 

(SO2), nitrogen oxide (NOx), particulate matter less than 

or equal to 10 microns or 2.5 microns in diameter (PM10

and PM2.5), ozone (O3), carbon monoxide (CO), and 

lead (Pb) (Fig. 4). The NAAQS were developed by the 

Environmental Protection Agency for a specific set of 

“criteria” pollutants to protect public health and welfare. 

NAAQS are defined as the amount of a criteria pollutant 

above which detrimental effects to human health or 

welfare may result. The NAAQS are set at conservative 

levels, with the intent of protecting even the most 

sensitive members of the public including children and 

people with asthma and cardiovascular disease (http://

www.epa.gov/air/criteria.html). Nevertheless, states can 

Figure 1.—Distribution of forests in the eastern United States.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 201

Figure 2.—Population density in the eastern United States.

Figure 3.—Locations of Interstate Highways overlaying population density in the eastern United 

States.
2	 Fire 02 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

adopt standards even more stringent than the federal 

standards. If an area consistently violates one of the 

NAAQS, that area will be federally designated as a “nonattainment” area for the specific criteria pollutant(s). 

States are required to demonstrate to the public and 

the EPA how emissions will be controlled in order 

for the non-attainment area to again comply with the 

NAAQS. These demonstrations are included in a State 

Implementation Plan (SIP), which may include methods 

to control emissions from prescribed fire.

6. Where/What are the Class I areas?

Mandatory federal Class I Areas were designated by the 

Clean Air Act amendments (CAAA) of August 7, 1977, 

and include national parks greater than 6,000 acres and 

national wilderness areas and national memorial parks 

greater than 5,000 acres which existed at that time (42

U.S.C. 7401 et seq). Resources such as soils, streams 

and visibility in these Class I Areas (called Air Quality 

Related Values) were given an additional measure of 

protection under the CAAA. Although the majority of 

Class I areas are in the west, several are scattered across 

eastern oak forests (Fig. 5). The eastern Class I areas are 

often located in, or near, densely populated areas, and 

receive extensive visitor use.

Regional Haze Rules address regional emissions that 

affect visibility in Class I areas. Recognizing that regional 

haze results from many sources of pollution over a 

large geographic area, these regulations require states 

(and tribes who choose to participate) to review how 

emissions generated within the State affect visibility 

at “Class I” areas. The EPA has encouraged states and 

tribes across the U.S. to address visibility impairment 

from a regional perspective and as a result the Regional 

Planning Organizations (RPO’s) (Fig. 6) were formed. 

These organizations are evaluating available technical 

information to better understand how the sources within 

their states and Tribal lands impact Class I areas across 

the country. This technical information can then be used 

by the states and tribes to develop regional strategies to 

reduce emissions of pollutants leading to regional haze 

(from: http://www.epa.gov/air/visibility/regional.html ).

Figure 4.—Locations of federally designated Ozone and PM2.5 non-attainment areas overlaying 

interstate highway locations and population density in the eastern United States.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 203

Figure 6.—Names and member states of the five Regional 

Planning Organizations

Figure 5.—Locations of Class I areas overlaying population density in the eastern United States.

States are required by the rule to analyze a pathway 

that takes the Class I areas from currently impaired 

conditions to “natural conditions” in 60 years. The term, 

”natural visibility conditions” is meant to “represent the 

long-term degree of visibility that is estimated to exist in 

a given mandatory Federal Class I area in the absence of 

human-caused impairment” (U.S. EPA 2003; pg. 1-1). 

States must show they are making “reasonable progress” 

toward achieving this goal. The Regional Haze Rule sets 

up the period of 2000 to 2004 as the baseline visibility 

conditions and “reasonable progress” will be tracked for 

that period onward. States currently are coordinating 

through the RPOs to determine the necessary level 

of reductions in haze forming emissions to achieve 

this. State Implementation Plans (SIPs) will need to 

be developed to outline strategies and schedules for 

emission reductions. Prescribed fire emissions are to be 

included in the suite of emission sources that potentially 

impact visibility. Most states will need to have a SIP in 

place by 2008 to cover a period of 10-years. While this 

program is aimed at Class I areas, visibility will improve 

throughout the country by reducing regional emissions.

Because the pollutants that lead to regional haze can 

originate from sources located across broad geographic 

areas, the EPA has encouraged the states and tribes across 

the U.S. to address visibility impairment from a regional 

perspective. Therefore, Regional Planning Organizations 

(RPO’s) (Fig. 6) have been formed to address regional 

haze and related issues. These organizations are 
2	 Fire 04 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

evaluating available technical information to better 

understand how the sources within their states and 

Tribal lands impact Class I areas across the country. This 

technical information can then be used by the states and 

tribes to develop regional strategies to reduce emissions 

of pollutants leading to regional haze (from: http://www.

epa.gov/air/visibility/regional.html).

5. Where and when does burning typically occur?

Figure 7 shows the counties in the eastern United States 

that reported federal prescribed burn activity in 2004 

( http://www.nationalatlas.gov/natlas/Natlasstart.asp). 

In addition to the locations indicated on this map, 

additional prescribed burn activity undertaken by state 

and local organizations occurs throughout the region.

4. Where does smoke typically go?

Smoke from prescribed burns has local, regional, and 

national impacts. Local impacts, such as those to sensitive 

receptors (homes, schools, businesses) directly downwind 

of a burn site are routinely addressed in burn plans. 

Regional impacts are sometimes included in burn plans, 

as some plans account for population centers away from 

the burn site. Additionally, there is evidence that smoke 

from prescribed burns can impact specific sites hundreds 

of miles away from the burn location, contributing 

to impacts more on a national scale. Figure 8 shows 

Figure 7.—Counties reporting prescribed fire activity in 2004 overlaying locations of oak forests 

in the eastern United States.

Figure 8.—48-hour back trajectories for 20% best (blue) and 

20% worst (red) haze days in the Shenandoah National Park, 

1997-1999 (from Regional Haze and Visibility in the Upper 

Midwest. Midwest Regional Planning Organization, (http://

www.ladco.org/tech/monitoring/docs_gifs/rhreport.pdf).
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 205

a 48-hour trajectory analysis for the best and worst 

visibility conditions measured at Shenandoah National 

Park from 1997-1999 (Midwest RPO, 2001). This 

analysis indicates that the source air a given location, 

such as a sensitive receptor or a national park can come 

from locations all across the eastern United States. The 

implication here is that any prescribed burn activity 

has the potential to impact air quality and visibility 

conditions on a regional and national scale. As states 

implement smoke management programs designed to 

monitor and, if necessary, alter burn programs to control 

air quality at specific sites, prescribed burn planning will 

become increasingly dependent upon modeling tools 

that assess the potential for smoke to influence regional 

and national air quality.

3. Where is burning expected to increase/decrease?

The USDA Forest Service Region 9, which stretches 

from Minnesota south to Missouri and east to the 

Atlantic Coast, treated (using prescribed fire and other 

fuel treatments) approximately 96,473 acres in FY 

2005 (USDA Forest Service 2006). Nationally, the 

USDA Forest Service in 2005 had a target of treating 

2.5 million acres of hazardous fuels through a variety of 

prescribed fire and other vegetation treatments. In FY 

2007 the target is approximately 3.2 million acres of 

hazardous fuels treatments. Each state also has treatment 

plans that include prescribed burning as an activity. 

Prescribed burning in mixed oak forests, and throughout 

the United States, is expected to increase in the near 

future.

2. Where are there coordinated Smoke Management 

Programs?

There is a wide variety in the type or existence of state 

Smoke Management Programs (SMPs). Florida has 

perhaps the best-established SMP in the East, with daily 

burning coordinated through a central clearinghouse in 

the Florida Department of Forestry. South Carolina and 

Minnesota likewise have SMPs in place. But many other 

states throughout the region are in the early stages of 

developing SMPs or have nothing at all (Fig. 9). As these 

programs become established and smoke management 

regulations are more clearly defined, the impact of smoke 

Figure 9.—Status of efforts to design and implement state Smoke Management Programs 

overlaying counties reporting prescribed burn activity in 2004 in the eastern United States. 
2	 Fire 06 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

management on prescribed burn planning and execution 

will become more apparent.

1. What does all this mean to fire and smoke managers?

When all of the elements listed above are considered 

together, it becomes clear that smoke management in 

the Oak forests can be extraordinarily complex. This 

burn activity takes place in a part of the country with the 

greatest population (compared to the west) and in states 

with existing air pollution problems and an extensive 

transportation system. Additionally, forest managers are 

hoping to increase the number and size of prescribed 

burns in the near future. Finally, coordinated SMPs 

designed to account for the cumulative effects of burning 

are few and far between in the region.

SMOKE MANAGEMENT TOOLS

There are many smoke management tools available to 

fire and smoke managers to aid in planning for a burn. 

The tools indicate how readily smoke will disperse from 

the fire area, and where the smoke will go in the ensuing 

hours. The National Weather Service provides forecasts 

of the Ventilation Index (VI) (Hardy et al. 2002; 

and others) in routine fire weather forecasts and spot 

forecasts requested by fire managers. The VI provides an 

assessment of the extent to which smoke produced by a 

fire will be effectively transported (ventilated) from the 

fire area. However, while the VI indicates how readily 

smoke will disperse, it does not indicate where the smoke 

will go once it leaves the burn site. A smoke dispersion 

model is necessary to determine where smoke is likely to 

go and what the pollution concentrations (mainly PM 

2.5) will be downwind of the burn.

The model that a fire or smoke manager employs 

depends strongly on what questions need to be answered. 

For questions concerning smoke concentrations within 

a few miles of a fire, dispersion models driven by single 

observations take at or near the fire location are often 

sufficient. VSMOKE (Lavdas 1996) is arguably the most 

advanced model that is currently in operational use in 

the eastern United States. VSMOKE is designed to help 

smoke and fire managers predict the smoke and dry 

weather visibility impact of a single fire at downwind 

locations. VSMOKE contains a plume rise model, a 

steady-state trajectory model, and other components 

that provide predictions of concentrations, visibility 

and other factors. Additional examples of dispersion 

models in use include: 1) the EPA’s Industrial Source 

Complex (ISC3) model, which is used for predicting 

dispersion of pollutants from industrial facilities mainly 

for permitting purposes (U.S. EPA 1995); 2) the Smoke 

Impact Spreadsheet (SIS), a simple planning puff 

model for calculating particulate matter emissions and 

concentrations downwind of wildland fires (Wickman 

and Acheson, 2005), and 3) The Simple Approach 

Smoke Estimation Model (SASEM), which predicts 

ground level particulate matter and visibility impacts 

from single sources in relatively flat terrain (Sestak and 

Riebau, 1988). All of these models can be used to assess 

smoke concentrations immediately downwind of a fire 

on time scales of a few hours. Additional details about 

the applications and relative merits of these and other 

smoke dispersion models can be found in Breyfogle and 

Ferguson (1996).

The Smoke Management Team at the USDA Forest 

Service’s Southern Research Station in Athens, GA is 

developing more advanced smoke transport models, 

the so-called Planned Burn (PB) models (Achtemeier, 

2001). PB smoke movement and dispersion models are 

designed to address the following questions: 1) if I burn 

tomorrow, will I smoke up a road?; and 2) I just burned, 

where is my smoke going to be tonight? The PB models 

address the problem of terrain and landscape features 

that can cause smoke plumes to diverge, split, merge, 

and evolve over complex local topography. The models 

predict smoke movement as a mixture of independent 

particles similar to smoke actually flowing downwind 

from a burn site. Smoke that becomes trapped in 

a localized area gradually dissipates over time. The 

“bleedoff” of smoke is simulated by allowing the model 

to occasionally create new smoke particles from existing 

particles that can then disperse over time (Achtemeier, 

2001). One of the major advantages of the PB modeling 

framework is that it is designed to incorporate highly 

detailed gridded weather information both at the ground 

and aloft. This detailed weather information allows the 

model to account for temporally and spatially varying 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 207

wind and stability conditions, and to predict how surface 

features (terrain, coastlines, etc) impact the smoke 

plumes as they evolve and disperse.

The first PB model to be developed was PB-Piedmont, 

which was designed to simulate smoke transport in 

the complex terrain of the southeastern United States 

Piedmont region. Two newer incarnations of the PB 

models have since been developed. PB-Coastal Plain 

incorporates land use data and land-water information, 

along with small variations in elevation, to model smoke 

movement over the lower Coastal Plain. Finally, PBMountain is designed to simulate smoke transport in the 

mountains of the Southeast. The Eastern Area Modeling 

Consortium (EAMC) is working to implement one or 

more PB models for the northeastern United States and 

eastern oak forests, and to make this smoke prediction 

tool available to fire and smoke managers across the 

region. Figure 10 shows an example of a one-time use 

of the PB-Piedmont model to simulate smoke from 

burning timber in Ohio.

The smoke dispersion models outlined above are 

generally applicable only to smoke management 

concerns in the immediate vicinity of a fire. None of the 

models are designed to address how smoke from fires 

could affect regional air quality. However, in Section 

2 we indicated that state smoke monitoring programs 

designed to address regional air quality issues will 

soon begin to impact smoke management decisions 

throughout the Eastern United States. Currently, there 

is no well-established tool that can help fire and smoke 

managers make regional smoke management decisions. 

However, researchers in the USDA Forest Service’s 

Pacific Northwest Research Station, in cooperation with 

universities and other research facilities, have developed 

the BlueSky Smoke Modeling Framework (O’Neill, 

2003) to address this need.

BlueSky is a modeling framework that brings the latest 

state-of-the-science fuels, fire, smoke, and weather 

models into one centralized processing system. It is 

designed to provide fire operations and air quality 

Figure 10.—PB-Piedmont simulation of smoke from burning a pile of Christmas trees at a 

refuse site near Lewisburg, OH in the Twin Creek Valley between 8:00 AM and 3:00 PM 

on 7 February 2000. Smoke from this fire, in addition to local dense fog, was implicated as 

a factor in a vehicle pileup on I-70 at approximately 10:55 PM on 7 February. 
2	 Fire 08 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

management communities easy access to 

sophisticated emission, dispersion and weather 

prediction models and model output. The 

modeling framework predicts the cumulative 

impacts of smoke from prescribed fire and 

wildfire on regional concentrations of PM2.5 and 

other particulates (Fig. 11).

The BlueSky system was designed as a tool to 

aide land managers in making go/no-go/go-slow 

decisions relative to smoke management from 

prescribed burns. The system produces hourly 

PM2.5 predictions based on input from the 

user and 209 wildfire reports. By employing 

centralized collection and processing of model 

data, the user does not need to download data and 

learn to use complex modeling systems. It also 

allows for analysis of multiple burns and wildfires 

so that air quality managers can see the combined 

impacts of prescribed burns within shared airsheds. 

BlueSky output is posted to the web daily for easy access 

by burners, air resource specialists, and the public.

Sample BlueSky output for the Pacific Northwest United 

States and publications are available at http://www.fs.fed.

us/bluesky. The Eastern Area Modeling Consortium 

(EAMC) maintains BlueSky for the northeastern United 

States. The user interface for submitting a prescribed 

burn to the EAMC implementation of BlueSky can 

be found at http://www.ncrs.fs.fed.us/eamc/products/

bluesky/.

CONCLUSION

This paper has outlined the pertinent smoke 

management questions that fire and smoke managers 

in the Central Hardwoods will face in the near future. 

As prescribed fire becomes a more common practice in 

ecosystem management, the potential for the resultant 

smoke to impact human health and public safety will 

have a correspondingly greater impact on burn plans 

and go/no-go/go-slow decisions on the day of the burn. 

We have outlined what we see as the main questions 

that land managers in eastern oak forests should be 

aware of as they develop forest management plans, as 

states implement smoke management plans, and as 

federal regulations that could affect smoke management 

decisions are being established. Additionally, we have 

provided an introduction to the existing tools and some 

of the new tools that are becoming available to aid in 

smoke management decisions. Clearly, prescribed burn 

planning in eastern oak forests will continue to become 

more complex as additional regulations are implemented 

and sensitivities both in the physical and social aspects 

of the problem become apparent. By remaining aware of 

the air quality regulations affecting prescribed burning 

programs and utilizing the tools available to assist them 

in meeting those regulations, land managers can be fully 

prepared for the changes that are certain to come.

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USDA Forest Service 2006. Fiscal Year 2007 President’s 

Budget - Budget Justification. 593 p. Specific 

citation on pp. 9-16 to 9-20.

Ward, J.S.; Brose P.H. 2004. Mortality, survival, 

and growth of individual stems after prescribed 

burning in recent hardwood clearcuts. In: Yaussy 

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Proceedings, 14th central hardwood forest conference; 

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Square, PA: U.S. Department of Agriculture, Forest 

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21	 Fire 0 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

RESISTANCE OF EASTERN HARDWOOD STEMS TO FIRE INJURY 

AND DAMAGE

Kevin T. Smith1† and Elaine Kennedy Sutherland2

Abstract.—This paper reviews the protective features and defensive responses of eastern 

hardwood species exposed to fire. Trees survive fire through protective features such as thick 

bark and the induced defenses of compartmentalization. Dissection of trees exposed to 

prescribed fire in an oak forest in southern Ohio highlights the need to distinguish between bark 

scorch, stem injury, and damage in evaluating the effects of fire on surviving overstory trees.

may be readily understood in terms of energy budgets. 

A budget reconciles outlays among various possible 

expenditures within the limits of income. An optimal 

budget strategy meets the needs of necessary expenditure 

with minimal surplus, leaving more resources available 

for other categories.

As green plants, trees capture and convert solar energy 

into chemical energy that is then dynamically allocated 

to meet the changing needs of the tree, including that 

of stored energy for future needs. Although strategies 

for allocation vary among species and settings, all trees 

need energy for growth, reproduction, protection, 

defense, and maintenance of the living system. One of 

the central dilemmas addressed by these strategies is 

the tradeoff between the allocation of energy to growth 

or to protection and defense (Loehle 1988; Herms 

and Mattson 1992). Height growth and a spreading 

crown architecture promotes energy capture under 

favorable conditions. Increased stem growth can enhance 

structural stability and supporter taller, broader crowns. 

On the other hand, investments such as thick bark and 

a heartwood that is enriched with wood-preserving 

chemicals can increase the probability of survival after 

injury.

One yardstick for describing the strategy for energy 

allocation is allometrics, the relative biomass distribution 

among various plant parts such as roots, foliage, wood, 

etc. Effective strategies promote while less effective 

strategies reduce the probability of survival. Tree species 

undergoing competition can have different strategies 

for energy allocation. These strategies vary among 

tree species and may also shift as trees increase in size 

(Jackson, Adams, and Jackson 1999). Trees survive fire 

BACKGROUND

Fires ignited by humans or by lightning are fundamental 

influences on eastern oak forests. In the absence of 

suppression efforts, these fires are generally frequent, 

low in intensity, and patchy in extent resulting in 

variable exposure of trees to fire (Wendell and Smith 

1986; Shumway, Abrams, and Ruffner 2001; Guyette 

and Spetich 2003). The eastern oak forests of today 

frequently contain an overstory that is comparatively 

fire-tolerant and a dense understory of more firesensitive and shade-tolerant species (Abrams 1990; 

Yaussy et al. 2003). In ecological terms, the current 

overstory is the likely result of regeneration or release of 

fire-tolerant trees prior to the onset of fire suppression. 

The current understory is dominated by fire-sensitive 

species that compete favorably with fire-tolerant species 

in the absence of fire. Under a scenario of continued 

fire suppression, the frequency of fire-sensitive species 

will likely increase in the canopy and remain high in 

the understory. The lower bole is potentially the most 

vulnerable part of a tree exposed to surface fires. How 

do some trees tolerate fire while others are sensitive? 

This review examines the basic survival mechanisms 

that reduce injury and damage to trees exposed to fire in 

eastern oak forests.

ENERGY BUDGETS AND TRADEOFFS

Although difficult to quantify, the concept of 

competitive advantage in the presence or absence of fire 

1

USDA Forest Service, Northern Research Station, 271 Mast 

Rd., Durham, NH 03824; 2Rocky Mountain Research 

Station, Forestry Sciences Laboratory, 800 East Beckwith, 

Missoula, MT 59801; †Corresponding author, 603-868-7624, 

email: ktsmith@fs.fed.us.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 211

exposure through the (1) constitutive, inborn allocation 

of energy for protective features that prevent injury 

and (2) induced allocation of energy for defense that 

maintains and restores tree health, particularly the 

continuity of the vascular cambium.

FIRE BEHAVIOR AND THE POTENTIAL 

FOR TREE INJURY

Patterns of fire injury reflect patterns of fire behavior. 

Most descriptions of fire behavior and its consequences 

have been based on observations of western conifers 

(Miller 2001) although some work has been done 

in eastern oak forests (Bova and Dickinson 2005). 

Although crown involvement of overstory trees may 

occur, eastern oak forests are most often subjected to 

surface fires fueled by fine or coarse woody debris and 

low-growing living plants. The direction and rate of 

movement of a forest fire depends on fuel quality and 

quantity, topography, and atmospheric conditions, 

including the prevailing wind direction.

Following ignition, a downwind-burning fire presents 

a fast-moving front with flames that typically burn the 

most readily combustible fuel and leave behind much 

charred and incompletely burned fuel. The upwind or 

backing fire is a much more slowly moving front that 

burns fuel more completely. The severity of tree injury is 

due to both fire intensity (amount of heat generated per 

length of front) and the duration of the period in which 

the flames are close to the stem.

As the flames of the heading fire pass a tree stem, flame 

height increases on the leeward side of the stem (Gill 

1974). This increase in height of the standing leeward 

flame is accompanied by increased residence time and 

greater stem heating (Gutsell and Johnson 1996). The 

same process increases heating on the upslope side of 

trees on uneven terrain as flames and the air entrained 

into the flames flows past tree stems.

CONSTITUTIVE PROTECTION

Activity by all of the meristematic tissues (shoot and root 

tips, vascular cambium, bark or cork cambium) results 

in compartmentation, the arrangement of tree cells into 

various tissues and organs (e.g. roots, foliage, branches, 

sapwood, heartwood) with specific roles in maintaining 

tree function. The unique characteristic of woody 

plants is the dominant role of the vascular cambium in 

tree survival and growth. The vascular cambium is the 

continuous sheath of small, fragile, thin-walled cells 

located immediately outside of the most recently formed 

wood and to the inside of the phloem or inner bark. The 

vascular cambium is the “new cell generator” for both 

xylem (that matures into wood) and phloem (sometimes 

referred to as inner bark, the transport tissue for most of 

the sugar and other biochemicals essential for tree life). 

An element of compartmentation is the annual growth 

layer or increment of wood in the boles, branches, and 

woody roots of temperate zone trees that appear in crosssection as tree rings. Essentially, a new sheath is formed 

every year over the core of the tree grown in previous 

years.

As an integrated system, all parts of the tree are essential 

for healthy functioning. However, we emphasize the 

survival of the vascular cambium in that it is more 

difficult to restore or replace than other plant tissues 

or organs (e.g. roots, foliage, branches). Also because 

of its anatomical position, death of the vascular 

cambium (sometimes referred to as “cambial necrosis”) 

also indicates the death of the phloem located to the 

outside and death of the sapwood located to the inside 

of the vascular cambium. In surviving trees, fire kills 

the vascular cambium and associated tissues by heat 

conducted through intact bark, rather than from direct 

combustion. This heat transfer can be modeled using 

a variety of approaches (Gutsell and Johnson 1996; 

Dickinson and Johnson 2001; Jones et al. 2004).

The vascular cambium of oaks is protected by a thick 

layer of bark. Bark is a poor conductor of heat and 

serves to insulate the vascular cambium from the heat 

of forest fire. Bark thickness has been used as the most 

obvious and readily measured protective feature that 

varies among tree species (e.g., Nelson, Sims, and Abell 

1933; Harmon 1984). However, other bark qualities, 

such as texture and the effect of developmental stage on 

bark thickness, also affects the effectiveness of bark in 

protecting the vascular cambium (Gignoux, Clobert, and 

Menaut 1997).
212 	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Traditionally, 60°C (140°F) has been used as the critical 

threshold temperature that kills the cells of the vascular 

cambium. This injury is termed cambial necrosis. 

However, there may be differences in the rate of cambial 

cell death at elevated temperatures among tree species 

and season of exposure (Dickinson and Johnson 2004). 

Cambial cell death also is affected by the duration 

of exposure to elevated temperatures and the rate of 

temperature increase (Dickinson, Joliff, and Bova 2004). 

INDUCED DEFENSE

Cambial necrosis provides access for the infection 

of wood by fungal decay pathogens and associated 

organisms. Injury and subsequent infection induce 

a cascade of responses within the constitutively 

compartmented portions of the tree. This process is 

collectively termed compartmentalization (Shigo 1984). 

Compartmentalization resists the spread of infection 

in wood. Compartmentalization is a boundary-setting 

process that consists of two parts. The first part of 

compartmentalization occurs in wood present at the time 

of injury. As living cells are injured, plugs form in the 

water-conducting cells that resist the spread of further 

cell death in the sapwood by aeration and desiccation. 

The plugging is accomplished both through the 

triggering of formation of specialized structures (tyloses) 

to block the axial water-conducting system as well as 

shifts in metabolism to produce waxes, gums, and resins 

that also restrict dessication and the entry of air into the 

xylem (Shigo 1984; Pearce 1996).

These shifts in metabolism of wood cells, as well as 

subsequent infection by fungi, tend to discolor or darken 

the affected wood. Because of wood anatomy, the spread 

of cell death and infection tends to occur in columns 

within the length of the stem. Boundaries are formed by 

living wood cells that limit the spread of wood cell death 

and infection to the smallest possible volume. These 

boundaries that resist the spread of the columns into 

healthy wood are referred to as reaction zones or column 

boundary layers. In oaks and other broadleaved trees, 

these layers frequently contain waterproofing materials 

similar to those found in bark and phenolic compounds 

such those found in heartwood and bark.

The second part of compartmentalization occurs in 

wood formed after injury. The vascular cambium 

produces an anatomically distinct barrier zone to both 

sides and above and below the injury. The barrier zone 

is frequently visible to the unaided eye as a darkened 

line within or between annual growth ring(s). The 

barrier zone is most obvious near the wound, but may 

extend around the tree circumference at some distance 

away from the injury. The position of the barrier zone 

within the growth ring allows the estimation of the 

timing of the fire within or between the growing seasons. 

In the absence of additional wounds that breach the 

barrier zone, wood decay will tend to occur within 

the compartment described by the barrier zone. The 

compartmentalization of infection and decay enables the 

vascular cambium to continue to divide and to move 

outward and away from the infection.

In an otherwise healthy, vigorous tree, the fire injury 

stimulates cell divisions of the vascular cambium at 

the edge of the area of cambial necrosis or fire scar, 

resulting in locally wide growth rings. This local growth 

stimulation serves to both hasten wound closure and to 

provide additional strength to the loadbearing structure 

of the tree. These thickened woundwood ribs serve to 

efficiently distribute the mechanical loading of the stem, 

branch, or root and to reduce the risk of structural failure 

(Mattheck 1998).

TREE DISSECTIONS AND 

QUALITATIVE ANALYSIS

We dissected eastern oak and associated trees previously 

exposed to prescribed fire at the Raccoon Ecological 

Management Area in southeastern Ohio (Yaussy, 

Hutchinson, and Sutherland 2003; Hutchinson, 

Sutherland, and Yaussy 2005). Our qualitative analysis 

(Smith and Sutherland 1999, 2001; Sutherland and 

Smith 2000) indicated that trees with scorched bark 

generally occurred in scattered clusters. Individual trees 

located near discrete fuel sources, such as snags and logs, 

were also scorched. Dissections showed that the vascular 

cambium of some scorched trees, particularly those with 

thick bark, was not injured by the prescribed fires. Thinbarked red maples were more frequently and severely 

injured than oaks and hickories. In oaks and hickory 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 213

that were slightly injured, cambial necrosis was localized 

to the areas beneath deep furrows in the bark, where the 

insulation provided by the bark was least.

Two years after fire injury, few if any overstory trees 

were killed by the fire. Most fire scars were concealed 

by intact bark. As with the pattern of bark scorch, these 

basal fire scars were triangular in shape (Smith and 

Sutherland 1999). This shape has been attributed to the 

triangular pattern of heat intensities within and around 

a flame (Gill 1974). However, basal wounds not caused 

by fire frequently form triangular scars due to increased 

dieback of the vascular cambium along the stem axis. 

Observations of trees exposed to two prescribed fires 

verify that the scorching or charring of wood in the open 

wound face resulted from exposure to a later fire and not 

to the fire that caused the initial scar.

Tree dissections showed that fire scars were often 

associated with bark fissures, particularly in tree species 

with thick bark (Fig. 1A, B). The relatively thin bark 

in the fissure provides less protective insulation to the 

vascular cambium beneath the fissure (Guyette and 

Stambaugh 2004; Smith and Sutherland 1999). The 

surviving vascular cambium adjacent to the fire scar 

generally produced xylem cells at an accelerated rate, 

resulting in a fold or rib of woundwood at the scar 

margin (Fig. 1A, B). This enhanced production of wood 

was seen as wider growth rings adjacent to the wound. 

This localized stimulation in growth tends to close over 

the exposed wood more quickly than at normal growth 

rates. Successful wound closure reduces the access of 

exposed wood and associated tissues to infection and 

allows for the potential restoration of the vascular 

cambium around the stem circumference.

The formation of woundwood ribs also contributes to 

the structural stability of the standing tree (Fig. 2A, B). 

In engineering terms, most of the mass of the aboveground portion of the tree is supported by the outermost 

portion of the roughly cylindrical stem (Mattheck 1998). 

Stem growth tends to uniformly distribute the loading 

Figure 1.—Cross- section of pignut hickory 

(Carya glabra) with small fire scars covered 

by intact bark, dissected in the third year after 

fire injury. A. Cambial necrosis (white arrows) 

associated with bark furrows. B. The column 

of wound-initiated discoloration (white arrows) 

is small and well-defined. Woundwood (ww) 

appears as locally wide annual rings. The small 

points of discoloration (black arrow) are likely 

the ends of small columns of discoloration 

initiated by a wound several centimeters above 

or below the plane of the section.
21	 Fire 4 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

around the stem circumference. Cambial necrosis 

locally interrupts the formation of the wood annual 

increment. As the stem continues to increase in girth, the 

physical loading becomes less uniformly distributed with 

additional stress applied at the margins of the wound 

(Fig. 2A, B). The tree woundwood ribs add support 

where the stress is increased. The curved orientation of 

wood cells at the inward-facing edge of the woundwood 

resists shearing stress parallel to the wound surface 

(Mattheck 1998).

The long-term effect of cambial injury caused by fire 

illustrates the effectiveness of compartmentalization in 

resisting the spread of infection and wood decay and 

contributing to tree survival. From the perspective of 

compartmentalization, a tree with a well-defined hollow 

or cavity can indicate a successful outcome (Fig. 3). 

The cavity is contained within a relatively small volume 

and surrounded by a continuous band of healthy and 

structurally sound wood. The vascular cambium is intact 

and not compromised.

IMPLICATIONS FOR MANAGEMENT

Evaluating the effects of fire on surviving trees, 

particularly low-intensity prescribed fire, requires an 

understanding of the distinctions and linkages among 

signs of a fire (e.g., bark scorch), stem injury (e.g., 

cambial necrosis, death of associated living phloem and 

sapwood, and resulting infection), internal discoloration 

and decay (limited by compartmentalization), and 

damage (loss of value). The mere presence of bark 

scorch does not necessarily indicate injury to the vital 

processes of a tree. Wound-initiated discoloration and 

decay can, within limits, be contained to small volumes 

within the living tree. The tree response to extensive 

injury and infection depends on induced defenses. The 

effectiveness of compartmentalization to resist the spread 

of infection depends on the genetic capacity of the tree 

for compartmentalization and the ability of the tree to 

use that capacity. That ability rests on the condition of 

the particular tree at the time of injury (Shigo 1986). 

The importance of pre-existing tree conditions for 

survival following fire injury was observed for eastern 

Figure 2..—Oak tree with a 4-year-old fire scar in 

southeastern Ohio. A. The scar is characterized by 

the exposed wood face that is almost completely 

covered by woundwood ribs. B. Simplified overlay 

diagram showing the mechanical loading of the 

stem (dotted lines). The decreased spacing of the 

load lines represents the increased load stress 

adjacent to the fire scar.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 215

oaks (Yaussy, Dickinson, and Bova 2004) as was 

previously seen for northern hardwoods exposed to 

storm injury (Shortle, Smith, and Dudzik 2003).

The distinction between injury and damage is 

especially important. “Damage” refers to a loss 

of value (Shigo 1986). Value is directly related 

to the specific goal of forest management. If 

prescribed fire shifts species frequency towards 

a more desirable stand composition, there may 

be significant injury to less-desirable species yet 

little or no damage in the sense of loss of value. 

Well-compartmentalized infections in the core 

of desirable tree species may result in a central 

column of wood decay or a cavity, but have 

little negative effect on the stem outside of that 

column, resulting in the potential for high wood 

quality for products or high habitat quality for 

wildlife. Depending on the severity of injury 

and the time required to restore continuity of 

the vascular cambium, fire can affect the value 

of overstory trees by decreasing the volume of 

comparatively more valuable heartwood both by 

increasing the volume of cull as discolored and 

decayed wood and by delaying the formation of 

heartwood (McGinness and Shigo 1975; Phelps 

and McGinness 1977). The degree of volume loss 

will depend on the extent of the injury and the 

effectiveness of compartmentalization.

CONCLUSIONS

Fire affects overstory trees in eastern oak forests 

in both obvious and hidden ways. Trees resist 

the occurrence of injury through constitutive 

protection and resist the spread of infection by 

induced defenses. Recognizing the patterns of fire 

effects can help guide prescriptions for the use of 

fire as a silvicultural tool and to understand the 

effects of wildfire. Bark scorch, tree injury, and 

damage are not equivalent terms. Useful evaluation 

of fire effects on and the implications for overstory 

trees requires an understanding of fire physics 

and tree physiology. The degree of damage sustained 

by fire injury depends on the specific goals of forest 

management.

Figure 3.—Cross-section of white oak (Quercus alba) illustrating 

the long-term response to cambial wounding. A. The tree 

was likely injured at the size and age indicated by the outer 

edge of the central cavity (small white arrows). Over the next 

approximately 40 years, woundwood ribs grew on either side of 

the exposed wound face, eventually restoring the continuity of 

the vascular cambium. Infection by wood decay fungi and their 

associates resulted in the complete decay of wood present at 

the time of injury. However, infection did not spread into wood 

formed after the injury. B. Changes in orientation of wood cells 

in woundwood ribs. Bark that lined portions of the inside of the 

cavity was produced on the outer surface of the ribs as they 

closed over the wound.
21	 Fire 6 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

ACKNOWLEDGMENT

We thank Ken Dudzik (USDA Forest Service, 

Northeastern Research Station) for two of the original 

photographs.

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21	 Fire 8 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Fire and Invasive Exotic Plant Species in Eastern Oak 

Communities: An Assessment of Current Knowledge

Cynthia D. Huebner1

Abstract.—Successful regeneration of oak-dominated communities in the Eastern United States historically 

requires disturbance such as fire, making them vulnerable to invasion by exotic plants. Little is currently 

known about the effects of fire on invasive plant species and the effects of invasive plant species on fire 

regimes of this region. Seventeen common eastern invaders were evaluated for their response to fire and 

potential to change current fire regimes. Twelve species are potentially controllable with repeated growingseason burns (decreasers); five may increase in abundance in response to fire (increasers). Most of the woody 

decreasers are also potential resisters of fire at maturity. The presence of a seedbank or an outside seed source 

(evaders) for all but one species and a positive germination response to post-fire conditions (e.g., higher 

soil temperature, nitrogen availability, and light) make it less likely that most eastern plant invaders can 

be controlled by fire alone. Shifts in fire regime in eastern oak communities are undocumented but may 

occur due to changes in community flammability after an invasion. Current fire models are inadequate for 

predicting fire behavior in these oak communities due to a lack of information on eastern native and exotic 

plant species fuels. Consequently, fire behavior predictions are best made on a site-by-site basis, especially for 

sites with multiple invaders composed of increasers and decreasers as well as fire promoters and inhibitors.

For this paper, the Eastern United States includes the 

Midwest, Great Lakes Region, Ozarks, Appalachian 

Mountains, Mid-Atlantic, New England, and Southeast. 

Within these regions are at least five community types 

with a dominant oak component that may be affected 

by fire and invasive exotic plant species: 1) mesic forests 

(mixed mesophytic), 2) dry forests (oak-hickory), 3) 

oak woodlands/savannas, 4) oak glades/barrens, and 

5) oak shrublands (scrub) (Jones et al. 1984; Andreas 

1989; Haney and Apfelbaum 1990; Grossman et al. 

1998). Definitions for woodland/savanna and glade/

barren overlap in the literature, but are separated here 

because glades/barrens have a more patchy distribution 

of trees than woodlands/savannas due to less available 

water or nutrients. Seventeen exotic plant species that 

are potential invaders of at least one or all of these five 

oak communities are evaluated for their response to and 

effects on fire (Table 1).

Fire and Exotic Plant Invasion History

Before 1850, most fires in the eastern region were surface 

fires set by native Americans. Fire intervals ranged from 

1 to 17 years, with shorter intervals in glades/barrens 

and woodlands/savannas than in forests. There was 

an increase in frequency between 1850 and 1930 due 

to European conversion of wilderness areas to farms. 

Between ~1850 and 1930 (and up to the 1950’s in the 

Introduction

The dependence of oak ecosystems on disturbance 

for regeneration (Loftis 1983; Cook 1998) can result 

in a vulnerability to invasion by exotic plant species, 

successful establishment of which often is linked to 

disturbance (Anderson 1999; Lonsdale 1999; Debinski 

and Holt 2000; Knapp and Canham 2000; Mack 

et al. 2000; Buckley et al. 2002). Disturbances that 

increase understory light (e.g., fire, harvesting, herbicide 

treatments or combinations of these) are effective in 

increasing oak regeneration (Dolan and Parker 2004; 

Miller et al. 2004; Rebbeck et al. 2004). If disturbance 

were the only important factor determining invasion 

success, oak systems requiring greater disturbance levels 

for maintenance would be the most likely to be invaded 

by exotic plant species. Factors other than disturbance, 

such as high resource availability (Richardson et al. 

1994; Burke and Grime 1996; Higgins et al. 1999; 

Lonsdale 1999; Stohlgren et al. 1999; Davis et al. 2000; 

Pysek et al. 2002; Thomson and Leishman 2005) and 

historic agricultural use (Dupouey et al. 2002, Ramovs 

and Roberts 2003) also can increase a community’s 

vulnerability to invasion.

1

USDA Forest Service, Northern Research Station, 180 

Canfield Street, Morgantown, WV 26505, 304-285-1501, 

email: chuebner@fs.fed.us.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 219

Table 1.—Seventeen exotic invasive species organized by habit type. Nomenclature in 

the table and text follows Gleason and Cronquist (1993) and the Integrated Taxonomic 

Information System (ITIS; 2005) http://www.itis.usda.gov/servlet/.

Herbaceous Plants

Canada thistle (Cirsium arvense (L.) Scop.)

Cogongrass (Imperata cylindrica (L.) Beauv.)

Garlic mustard (Alliaria petiolata (M. Bieb))

Japanese stiltgrass (Microstegium vimineum (Trin.) A. Camus)

Leafy spurge (Euphorbia esula L.)

Spotted knapweed (Centaurea biebersteinii DC.)

Vines

Japanese honeysuckle (Lonicera japonica Thunb.)

Kudzu (Pueraria montana (Lour.) Merr. var. lobata (Willd.) Maesen & Almeida)

Oriental bittersweet (Celastrus orbiculatus Thunb.)

Shrubs

Autumn olive (Elaeagnus umbellata Thunb.)

 (Russian olive – E. angustifolia was used as an example of EHOC value in Table 2 that may 

 correlate with autumn olive)

Bush honeysuckle (Lonicera L. spp.)

 Data in the text was presented specifically for Morrows bush honeysuckle (L. morrowii A. 

 Gray), Amur honeysuckle (L. maackii (Rupr.)), and Maxim and Bell’s bush honeysuckle (L. x 

 bella Zabel).

Common buckthorn (Rhamnus cathartica L.)

Japanese barberry (Berberis thunbergii DC.)

 Common barberry (B. vulgaris L.) was used as an example of EHOC value in Table 2 that may 

 correlate with Japanese barberry.

Multiflora rose (Rosa multiflora Thunb.)

Privet (Ligustrum L.spp.)

 Chinese privet (L. sinense) and common privet (L. vulgare L.) had specific research data 

 described in the text.

Trees

Norway maple (Acer platanoides L.)

Tree of heaven (Ailanthus altissima (Mill.) Swingle)

Ozarks) catastrophic wildfires increased after logging. 

Before this time, fire-rotation periods generally were less 

than 10 years, though this varied by community type 

and region. After 1930-50, fire suppression increased 

rotation periods to as long as 6,000 years for the Eastern 

United States (Dey 2002). Despite these changes in fire 

frequency and rotation, most fires in the East were and 

remain human caused and often are correlated with 

drought (Dey 2002; Muzika et al. 2005).

Recorded invasions of exotic plants and other exotic 

species since 1800 have increased at an accelerated rate, 

presumably due to increased intercontinental mobility 

(Liebhold et al. 1995). Similar increases in exotic plant 

species at a more local scale are evident from herbarium 

data (Huebner 2003). Although the correlation is 

circumstantial and weaker for the exotics, there is a 

noticeable increase in all species collected after the 

intensive logging and burning phase (1879-1920) in 
22	 Fire 0 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

West Virginia (Clarkson 1964). Thus, while fires in the 

Eastern United States have decreased since the 1950’s, 

exotic invasive plants have increased since the 1800’s, the 

tenuous connection being human intervention (Fig. 1).

Native Oak Communities’ 

Response to Fire

In general, oak species may benefit from fire because: 

1) intense fires followed by low-intensity fires open 

areas for encroachment by oak species and/or 2) lowintensity fires in forests open the subcanopy or canopy 

and reduce competition from shade-tolerant species 

(Lorimer et al. 1994; Miller et al. 2004). Hom (2003) 

proposed that shrub oak barrens, oak woodlands, and 

oak forests require fire-return intervals of 15 to 25, 20 to 

30, and 100 to 200 years, respectively, for community 

maintenance. These intervals are inversely proportional 

to each community’s likeliness to burn (Streng and 

Harcombe 1982).

Spring burns are often more successful at reducing 

non-oak woody vegetation with epigeal germination 

(cotyledons are above ground) and slow root-system 

development that contrast with oak species which 

generally have hypogeal germination (cotyledons are 

below ground) and relatively rapid root growth. Fall and 

winter burns have less effect on woody species, which 

have translocated much of their energy into belowground storage (VanLear and Brose 2002; Richburg et 

al. 2001). Repeated spring and fall burns may increase 

prairie forb cover in oak woodlands/savannas and oak 

glades/barrens by reducing competition of grass species, a 

possible management concern for threatened forbs, e.g., 

wild lupine (Lupinus perennis L.) (Pauly 1997).

Flammability and Fuels of Invasive 

Plant Species

Flammability has four components: 1) combustibility, 

2) ignitability, 3) consumability, and 4) sustainability 

all of which are influenced, on an individual plant basis, 

by plant moisture content, percent make-up of carbon, 

presence of volatile compounds, leaf thickness, and 

overall surface area-to-volume ratio (Behm et al. 2004). 

A few comparisons have been made on combustibility 

of eastern native and exotic plants using effective 

heat of combustion (EHOC; quantified with a cone 

Figure 1.—Time trend in percentage of the total number of species in herbarium collections 

from West Virginia for nine exotics (o), pteridophytes/gymnosperms (diamond), dicots (square), 

Asteraceae members (triangle), Solidago canadensis (x) (Huebner 2003) overlaid by total 

number of exotic insect and mite species recorded in the US (Liebhold et al. 1995) and the 

estimated fire rotation periods in the eastern hardwood forests (Dey 2002). The nine species 

include three bush honeysuckles (amur, morrow, and tatarica), garlic mustard, Japanese 

honeysuckle, Japanese stiltgrass, multiflora rose, oriental bittersweet, and tree of heaven. 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 221

calorimeter) (Dibble et al. 2003b). Compared to western 

fuels’ combustibility (14.5 to 21.6 MJ/kg), fuels of both 

native and nonnative species in the Eastern United 

States tend to have lower EHOC values (10.9 to 16.0 

MJ/kg). These studies also showed that nonnative species 

had lower EHOC values (10.0 to 14.2 MJ/kg) for 

combustibility than native species (11.7 to 16.0 MJ/kg; 

Table 2). The available EHOC data weakly suggest that 

eastern invasive plant species may tend to reduce fire 

occurrence; flammability data, using all components, 

on more species are needed (White et al. 2002). The 

ability to predict fire behavior in oak communities also 

requires knowledge of the flammability of the fuel bed. 

Surface fuels beds are composed of a range of categories, 

including litter, downed wood, shrubs, grasses, and forbs 

and are described by their average depth and percent 

cover and weighted averages of the 1) loading (lbs/ft2

), 2) 

surface-area-to-volume ratio, 3) heat content (BTU/lb), 

and 4) moisture content of each sub-class of fuel. Fuel 

sub-classes are determined by the general fuel category 

and the size of particles within a category and whether 

the material is dead or alive (Richburg et al. 2004). 

Community flammability is determined by multiple, 

interacting species, and, consequently, is more difficult to 

measure than flammability of individual species.

Species and community flammability, as well as current 

weather conditions, are all needed to predict fire 

behavior at a site. Models like BEHAVE’s NEWMDL 

program (allowing for custom fuel models) may 

improve predictive capabilities by incorporating data 

from eastern forests (Richburg et al. 2004). However, 

it is essential that the fuel bed characteristics of each 

community type be well-defined. For instance, in 

woodlands/savannas, the surface-to-volume ratio of fine 

fuels has a disproportionately large effect on fire spread, 

whereas in a closed hardwood stand, both the surfaceto-volume ratio and the fuelbed depth of fine fuels have 

a disproportionately large effect on the spread rate of 

fire (Ducey 2003). In a study with paired invaded and 

noninvaded mixed hardwood and hardwood stands, the 

invaded stands had less nonwoody litter, more 100-hr 

fuels, less duff depth, more graminoid and shrub cover, 

and less basal area than the noninvaded stands (Dibble 

et al. 2003a). Thus, while invasive plants may have low 

individual flammability, they may change a community’s 

flammability such that fire can burn and spread rapidly 

due to the presence of graminoids, or burn slower and 

more intensely in the presence of 100-hr fuels.

Exotic Plant Invaders’ Response to Fire

Invasive plant species responding to fire can be classified 

as: 1) evaders -- species with long-lived propagules stored 

in the soil (or a reliable outside seed source), 2) endurers

-- resprouters, or 3) resisters -- species that survive 

low-intensity fire due to certain adaptive characteristics. 

These can be classified secondarily as increasers -- likely 

to increase in abundance after a burn, maintainers

-- likely to maintain their current population size after 

a burn, or decreasers -- likely to decrease in abundance 

after a burn (Harrod and Reichard 2001). Each of the 17 

species evaluated has been categorized on the basis of the 

available literature (Table 2).

Several nonnative woody species, including autumn 

olive (Szafoni 1991a), common buckthorn (Solecki 

1997; Richburg et al. 2004); Chinese privet (Faulkner 

et al. 1989); Japanese barberry (Richburg et al. 2004); 

Japanese honeysuckle (Barden and Matthews 1980; 

Solecki 1997); kudzu (Radar et al. 1999); Morrow’s 

and Bell’s bush honeysuckle (Nyboer 1992; Luken 

and Shea 2000; Solecki 1997; Richburg et al. 2004); 

multiflora rose (Szafoni 1991b; Solecki 1997); Norway 

maple (Simpfendorfer 1989; Maissurow, 1941, based 

on similarities with sugar maple); and two herbaceous 

plants, garlic mustard (a biennial) (Nuzzo 1996; 

Schwartz and Heim 1996) and spotted knapweed (a 

perennial), can be reduced in community importance 

after repeated (annually consecutive for at least 2 to 5 

years) growing-season (spring to early summer) fires. 

The data for spotted knapweed, which can resprout 

after a burn like all of the above species (Sheley et al. 

1998), are unpublished2

. Japanese stiltgrass, an annual, 

can be expected to decrease in cover after a burn, given 

no seed source, though there is no direct evidence for 

this and resprouting is possible (Tu 2000; Gibson et al. 

2002). Though all 12 of these species are technically 

2

Zouhar, K. 2001. Centaurea maculosa. In: Fire Effects 

Information System. U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Research Station, Fire Sciences 

Laboratory. Available: http://www.fs.fed.us/database/feis/. 

Accession date: October 20, 2005.
222 	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Table 2.—EHOC values, colonizer type, response to fire, and possible effects on fire regimes 

documented for each exotic invasive plant species. Species are judged based on their response to 

repeated spring fires. na = no data; ? indicates no literature is available to support the possible 

categorization.

Species EHOC

(MJ/kg)

Colonizer type after fire Population response 

to fire

Promote or 

inhibit fire

Herbs

Canada thistle na Endurer Increaser Promote? 

Evader

Cogongrass na Endurer Increaser Promote?

Evader

(off-site seed only)

Garlic mustard na Evader Increaser Neither?

Decreaser

Japanese stiltgrass 11.14 Evader Increaser Promote?

Decreaser

Leafy spurge High (no value) Endurer Increaser Promote?

Evader

(but hot fires kill seed)

Spotted knapweed na Evader Increaser Inhibit?

Decreaser

Vines

Japanese honeysuckle 14.34 Evader Decreaser Promote?

(but poor seed 

production)

Maintainer (as ladder fuel) 

Inhibitor?

Resister

Kudzu na Evader Maintainer Inhibit?

Resister Decreaser Promote?

(older individuals) (younger plants) (as ladder fuel)

Oriental bittersweet 11.44 Endurer Increaser Inhibit? 

Evader Promote?

(off-site seed only) (as ladder fuel)

Shrubs

Autumn olive na (13.42) Evader Increaser Neither?

(Russian olive) (off-site seed, possible 

seedbank)

Decreaser

Bush honeysuckle na Evader Increaser Neither?

(off-site seed only) Decreaser

Common buckthorn na Evader Increaser Neither?

(off-site seed, possible 

seedbank)

Decreaser

Resister Maintainer

(older individuals)

continued
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 223

Japanese barberry na (14.04) Evader Increaser Neither?

(common barberry) (off-site seed source, 

possible seedbank)

Decreaser

Multiflora rose 12.30 Evader Increaser Neither?

Decreaser

Privet na Evader Increaser Neither?

(off-site seed only) Decreaser

Trees

Norway maple 11.11 Evader Increaser Inhibit?

(off-site seed; seedbank 

possible but unlikely)

Maintainer

Resister Decreaser

(older individuals)

Tree of heaven na Endurer Increaser Neither? 

Evader

(off-site seed only)

Table 2.—continued.

Species EHOC

(MJ/kg)

Colonizer type after fire Population 

response to fire

Promote or 

inhibit fire

endurers, they may be considered decreasers because 

of the evidence that their underground reserves and 

ability to sprout may be insufficient to withstand 

repeated growing-season burns. However, this evidence 

is not limited to oak-dominated communities; some 

studies were conducted in pine plantations (Japanese 

honeysuckle), prairies (spotted knapweed) and pastures 

(multiflora rose). Also, in the case of garlic mustard, there 

is evidence of increased garlic mustard abundance after 

repeated growing season burns in oak systems (Nuzzo 

et al. 1996; Luken and Shea 2000). The absence of a 

seedbank or outside seed source of all these species would 

ensure their status as decreasers or maintainers (Table 2).

Conversely, fires may be ineffective against large, mature 

shrubs of common buckthorn (Solecki 1997) and against 

large, woody individuals of any of these species. Similarly, 

kudzu can maintain a high water content due to its deep 

taproot so it may not experience topkill during a growing 

season burn (Winberry and Jones 1973). Tissue moisture 

content of Japanese honeysuckle (Slezak 1976) and 

Norway maple (Horvitz et al. 1998) may also be resistant 

to some fires. Thus, depending on age and size, autumn 

olive, common buckthorn, Japanese barberry, Japanese 

honeysuckle, kudzu, the bush honeysuckles, multiflora 

rose, privet, and Norway maple may be classified as 

resisters and maintainers.

Several species with deep and/or extensive underground 

root reserves, including oriental bittersweet (Dreyer et 

al. 1987), tree of heaven (Lepart and Debussche 1991), 

Canada thistle (Adams et al. 1987; Solecki 1997), 

leafy spurge (Cole 1991), and cogongrass (King and 

Grace 2000; Grace et al. 2001) may respond positively 

to repeated growing-season burns by increasing in 

abundance due to an ability to root sucker or sprout, 

keeping in mind that the research is not directly related 

to fire response (oriental bittersweet and tree of heaven) 

or took place in a pine plantation (cogongrass) or prairie/

grassland (leafy spurge and Canada thistle). These species 

can be considered potential endurers and increasers

after a fire (Table 2). Although evidence is stronger that 

Canada thistle responds positively to fire, some studies 

show it decreasing in abundance after summer or fall 

burns (Kirsch and Kruse 1973; Hogenbirk and Wein 

1991). Seedlings of cogongrass respond positively to a 
22	 Fire 4 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

burn and this grass is likely to respond to fire like other 

perennial warm-season grasses. However, its growth 

response to multiple fires in different seasons is unclear 

(King and Grace 2000; Grace et al. 2001).

Seed germination of kudzu often is promoted after a 

fire due to scarification (Takahashi and Kikuchi 1986; 

Susko et al. 2001). Garlic mustard (Hintz 1996; Byers 

and Quinn 1998), Japanese stiltgrass (Barden 1987; 

Anderson and Schwegman 1991; Anderson et al. 

2000; Glasgow and Matlack 2005), multiflora rose 

(Szafoni 1991b; Kaye et al. 1995; Leck and Leck 1998; 

Luginbuhl et al. 1999; Glasgow and Matlack 2005), 

spotted knapweed (Davis et al. 1993), Canada thistle 

(Kellman 1970; Turner et al. 1997), and leafy spurge 

(Wolters et al. 1994) all have seedbanks and may respond 

positively to fire (or, rather, the resulting removal of litter 

and increases in soil surface temperature and nutrients) 

with respect to on-site seed germination. Thus, these 

species are potential evaders and increasers (Table 2).

Seed banking for tree of heaven (Kowarik 1995, but see 

Kostel-Hughes and Young 1998), oriental bittersweet 

(Van Clef and Stiles 2001; Ellsworth et al. 2004), 

bush honeysuckles (Lonicera maackii (Rupr.) Maxim.; 

Ingold and Craycraft 1983), privet (Ligustrum sinense

Lour. and L. vulgare L.; Panetta 2000; Shelton and 

Cain 2002), cogongrass (Grace et al. 2001), and garlic 

mustard (Baskin and Baskin 1992) is minimal and most 

establishment by seed after a fire is by off-site dispersal. 

It is uncertain whether Japanese barberry, autumn olive 

(Katz and Shafroth 2003), common buckthorn, and 

Norway maple (Hong and Ellis 1990) have seedbanks. 

Tree of heaven seed is dispersed as far 200 m by wind 

(Graves 1990; Kota 2005); Norway maple also is 

wind dispersed but distances are not known. Oriental 

bittersweet, autumn olive, and multiflora rose have 

been dispersed as far as 500 m by starlings (LaFleur and 

Rubega 2005), while Japanese barberry seeds have been 

dispersed at least 80 m by birds (Silander and Klepeis 

1999). Bush honeysuckle and common buckthorn also 

are dispersed by birds but distances are not published. If 

outside seed sources exist, all of these species would be 

potential evaders and increasers. There is little evidence 

of Japanese honeysuckle having a seedbank and this 

species is a comparatively less prolific seed producer due 

to pollinator limitations (Larson et al. 2002) and low 

fruit viability (Haywood 1994). Thus, establishment by 

seed of Japanese honeysuckle after a fire may be unlikely. 

A species’ population response to fire may differ from its 

individual response. For instance, a patchy forest burn 

likely only removes portions of a large garlic mustard 

population. Because the population growth of this 

species is density dependent (Meekins and McCarthy 

2002), a large population of garlic mustard likely will 

show a positive growth response after a patchy burn due 

to a decrease in intraspecific competition.

Potential to Change Current Fire 

Regimes

There are no specific data showing shifting fire regimes 

in the Eastern United States for the 17 species evaluated. 

Leafy spurge may alter fire intensity because of the high 

oil content in its leaves (Davis 1990). However, fires 

with this species present may become hot enough to kill 

leafy spurge (and native species) seedbank seeds (Wolters 

et al. 1994). This could halt leafy spurge regeneration 

if sprouting were unlikely (Brooks et al. 2004). Canada 

thistle, Japanese stiltgrass, and cogongrass can build up 

fine, continuous fuels such that a rapidly moving, even 

spreading fire is likely should a burn occur (Barden 

1987; Hogenbirk and Wein 1995; Rice 2004). Such 

species may cause similar changes in fire regime as was 

found with the burn and regeneration cycle of cheat grass 

(Bromus tectorum L.; D’Antonio and Vitousek 1992; 

Brooks et al. 2004). The mesic habitats that Japanese 

stiltgrass tends to invade and the low flammability of this 

grass (compared to western species) may make such a 

regime change unlikely.

If a community is invaded by vines, such as oriental 

bittersweet, the vine may act as a ladder fuel that could 

result in a rapidly spreading canopy burn under ideal 

climatic conditions. Spotted knapweed tends to increase 

the patchiness of the grassland communities it invades, 

making it more difficult for such communities to carry 

a fire (Sheley et al. 1998). Also, if shrubs are invading 

a community that was not previously dominated by 

shrubs, there may be a subsequent decrease in fine fuels 

and a reduction in the community’s ability to carry 

a fire, though any fire that does occur may be more 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 225

intense. If the community is a savanna or barren, the 

latter may result in the loss of that community type over 

time. However, such resource-limited communities may 

not be invaded as easily. Japanese honeysuckle (Slezak 

1976) and Norway maple (Horvitz et al. 1998) may 

promote an increase in shade-tolerant species by shading 

out intolerant species, which, in turn, may reduce the 

likelihood of a burn. These species would be similar to 

the Brazilian pepper tree (Schinus terebinthifolius Raddi.), 

which retains high moisture in its leaves and litter 

and has reduced the fire frequency in once pyric pine 

rocklands (Gordon 1998).

Conclusion

If an invaded site is being managed for oak regeneration, 

the repeated burning required to control certain invasive 

species, e.g., bush honeysuckle and privet, may have a 

detrimental effect on oak, e.g., northern red oak (Quercus 

rubra L.), and other native species in the more mesic 

communities. However, such burns may be required to 

maintain certain xeric forests, woodland/savanna and 

glade/barren communities, though repeated burns might 

promote the invasion of other nonnative species, e.g., 

leafy spurge and Canada thistle.

We still lack adequate data with which to discern reliable 

patterns. The comparatively well-studied species, garlic 

mustard and Canada thistle, have variable responses to 

fire. As additional research is conducted on these 17 and 

other species, it is likely that the patterns discussed here 

will become more complex and variable. Such variability 

probably will result from differences in site conditions 

(community flammability, species flammability, resource 

availability, land-use history) before and after a burn 

(which are often unknown or difficult to measure) as 

well as each invasive plant’s population age structure 

and size just before a burn (both of which also are 

often unknown and difficult to measure). Moreover, 

many of these species occur together and joint effects 

on how a site will burn may differ from individual 

effects. Management of communities with multispecies 

invasions in which multiple increasers and decreasers as 

well as promoters and inhibiters might be present may 

be of such a complex nature that a site-specific approach 

always is required.

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Rx FIRE LAWS: TOOLS TO PROTECT FIRE: THE ‘ECOLOGICAL IMPERATIVE’

Dale Wade1†, Steven Miller2, Johnny Stowe3, James Brenner4

continued to learn new ways to improve their standard of 

living through the use of fire. Henry Lewis (1973) listed 

70 reasons Native Americans burned the land while 

Williams (2000) grouped their reasons for burning into 

11 categories. The pattern of occasional higher intensity, 

wind-driven fires and severe-drought fires that was 

superimposed on chronic lightning and Native American 

fire regime shaped and maintained vast southern prairies, 

savannas, open woodlands, and canebrakes, according to 

early European explorers. The journals of many of these 

explorers also mentioned numerous smoke columns 

and extensive smoke and haze that often lasted for days 

(e.g., deLaudonniere 1587). Into this environment 

came European—more than 75 percent with a pastoral 

background—(Owsley 1945) and African settlers. Both 

groups brought a knowledge of fire that they merged 

with that of the aboriginal American residents. Thus, the 

existing fire regime was expanded and reinforced.

The region’s endemic flora and fauna thus became 

even more intermingled with and dependent on fire 

(Martin and Sapsis 1992). Eldredge (1911) described 

the turn-of-the-century fire situation in north Florida: 

“The turpentine operator burns his woods and all other 

neighboring woods during the winter months, generally 

in December, January, or February. The cattleman sets 

fire during March, April, and May to such areas as the 

turpentine operator has left unburned. During the summer 

there are almost daily severe thunderstorms, and many 

forest fires are started by lightning. In the dry fall months 

hunters set fire to such “rough” places as may harbor game. 

It is only by chance that any area of unenclosed land escapes 

burning at least once in two years.” This pattern typified 

INTRODUCTION

We believethe Florida Prescribed Fire Act (Appendix A) 

is the best such statute in the country. If you participate 

in crafting a bill to safeguard the intentional use of fire 

in your area, we suggest you use the Florida act and 

accompanying rules as your starting point and modify 

them as necessary. That is the easy part; the real challenge 

is to get enough of the “right” people actively involved 

to sway public opinion and assure passage of your bill. 

The key to such an endeavor is an underlying faith that 

well-informed people will make well-informed choices. 

Because we have such faith, we weave our message and 

suggestions into a tapestry of fire and fire-use history in 

the hope that we will motivate you to action.

FIRE USE IN THE SOUTH

In the South, the ecological fire-return interval is shorter 

than almost anywhere else in the Nation, the result of 

both natural and anthropogenic fire. The incidence of 

cloud-to-ground lightning is higher in the South than in 

any other region of North America, and is responsible 

for many wildland fires. But anthropogenic fires are of at 

least equal importance. “By about 11,000 years ago, the 

Paleo-Indians and their fires had traversed all of the New 

World from Alaska to the tip of South America” (Stowe 

2004a). Over the ensuing centuries, Native Americans 

1

The Rx Fire Doctor, LLC, PO Box 491, Hayesville, NC 

28904-0491; 2

Division of Land Management, St. Johns River 

Water Management District, 4049 Reid Street, Palatka, FL 

32177; 3Heritage Preserves, South Carolina Department 

of Natural Resources,1000 Assembly Street, Columbia, SC 

29201; 4Florida Division of Forestry, 3125 Conner Boulevard, 

Tallahassee, FL 32399-1650; †Corresponding author, email: 

rxfire@ix.netcom.com.

Abstract.—The South is the birthplace of statutes and ordinances that both advocate and protect 

the cultural heritage of woods burning, which has been practiced in this region uninterrupted for 

more than 10,000 years. We present a brief overview of fire use in the South and discuss why most 

southern states recognized early on that periodic fire was necessary to sustain fire adapted ecosystems 

and passed laws to protect and facilitate the practice of controlled burning. We also provide examples 

of legislation promoting this ecological imperative and share ideas we have found helpful in getting 

the “right” people involved to assure passage of such legislation. Finally, we discuss constraints to 

prescription fire that need to be mitigated, as well as items to consider including in pro-fire legislation.
2	 Fire 34 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

the situation throughout the region, and although not 

everyone was in agreement with this ubiquitous use of 

fire, the survival of rural families depended on it (see 

Komarek 1981). So long as people accepted fire and its 

inevitability on nature’s terms, the precarious balance 

between humans and this natural force was maintained. 

Rural southerners realized that fire exclusion would tip 

this balance, destroying both ecosystems and attendant 

wildlife, while significantly increasing the likelihood 

that they would lose their homes and even their lives 

to uncontrollable wildfires that follow attempted fire 

exclusion.

When it became obvious by the late 1920’s that rural 

residents of the South were openly ignoring state and 

federal laws banning fire, the USDA Forest Service 

sent in the Dixie Crusaders (Schiff 1962; Jacobs 1978) 

to preach the benefits of fire exclusion and the evils of 

woods burning. Sociologists and psychologists such as 

John Shea (1940) also were sent with orders to “find the

inner-most reason why inhabitants of the forest lands of the 

South cling persistently to the custom of burning the woods” 

so that a more effective fire prevention campaign could 

be mounted and finally put a stop to “the annual fires 

that for more than a century have characterized the land 

and social economy of our southern states.” Shea (1940) 

thought the nearly unanimous explanation given by 

people he interviewed “that woods fires kill off snakes, 

boll weevil(s) and serve other economic ends are something 

more than mere ignorance. They are the defensive beliefs of a 

disadvantaged culture group.” History proved Shea wrong 

but the point is that it is only natural for outside experts 

as well as uneducated and misguided but otherwise 

well-meaning people to attempt to “correct” customs and 

actions that they do not believe are in the best interests 

of those who practice them. Such customs and actions 

may require adjustment periodically in accordance 

with John Dewey’s “reflective morality” (Frankena and 

Granrose 1974; Stowe 1997), but attempted “force 

feeding” usually promotes only rebellion.

When you have an objective that requires the help of 

people with different opinions, your agenda more than 

likely will be advanced if you keep an open mind, state 

your views clearly, listen and internalize what you are 

told, put yourself in their shoes, and, if you still disagree, 

craft your rebuttal from their point of view. This process 

is much more likely to yield a workable solution than 

if you “talk down” to people, or relentlessly harangue 

them with your message. More extensive discussions of 

southern fire history can be found in Komarek (1981), 

Pyne (1982), Wade and others (2000), Johnson and Hale 

(2002), and Stowe (2004a, b). Putz (2003) is a must 

read for anyone attempting to understand the culture of 

woods burning in the South; he provides a humorous 

but accurate account of why historic landscapes have 

been maintained in many areas of the South despite 

federal and state laws that once banned the practice 

of burning the woods. Future laws that ignore or ban 

local traditions and culture likely also will be ignored. 

On its website, the Florida Division of Forestry puts it 

succinctly: “Removing the option of controlled burning does 

not stop the burning, it just removes the control.”

Many people questioned the southern practice of 

woods burning and devised experiments to study this 

phenomenon, which led to the science of prescribed fire. 

These studies and observations, many by government 

land managers and by Herb Stoddard of the Tall Timbers 

Research Station (See Komarek 1981), moved the science 

forward until more acres are prescribe burned in the 

South today than in the rest of the nation combined. 

Wade and others (2000) estimated that more than 3 

million acres of forest land and 3 million acres of pasture 

in the South are treated with prescription fire each year, 

down about 2 million acres from the 1970’s (Wade and 

Lunsford 1989). Haines and Busby (2001) reported that 

slightly more than 4 million acres of southern pine were 

burned annually between 1985 and 1994. The 13 states 

in the Sourh comprise more than 534 million acres, 

most of which burned periodically a couple of centuries 

ago, so there is an enormous gap between the acreage 

that burned historically and the acreage that currently 

burns. This disparity is a major reason why the South 

typically has more wildfires than all other sections of the 

contiguous United States combined (more than 45,000 

per year). Although these fires tend to be smaller, the 

South often has more acreage subjected to wildfire as 

well, averaging nearly 1 million over the past 7 years 

(Gramley 2005).
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 235

THE NATURAL ROLE OF FIRE

The complexities of combustion aside, fire is nothing 

more than the rapid oxidation of organic matter, the 

same basic process as decomposition, albeit at a much 

faster rate. Wildland fire is a natural process that 

produces change—a basic premise underpinning the 

field of ecology. Whether these changes are viewed as 

desirable or not depends on their compatibility with 

human values, which typically have little do to with 

the natural scheme of things. Fire starts, facilitates, 

accelerates, decelerates, or stops the myriad of natural 

processes necessary to perpetuate fire-adapted ecosystems 

(see Wright and Heinselman 1973; Christensen 1977; 

Wade and others 1980). Sooner or later, fire is required 

to rejuvenate or maintain most terrestrial ecosystems and 

the various ancillary loops and cycles that keep them 

fully functional, though its return interval can range 

from months to centuries. In-depth discussions of the 

role of fire in North American ecosystems are found 

in DeBano and others (1998), and Brown and Smith 

(2000), both of which contain extensive references.

Fire has been shaping Earth’s landscape for millions 

of years, and has been augmented by anthropogenic 

ignitions for a long time (See Robinson 1988). Fire not 

only helped shape and maintain the terrestrial biota 

around us but it profoundly influenced the progress 

of humans. Fire was the first tool we learned to use 

on a landscape level, and our ability to harness this 

natural force sets us apart from all other animals. The 

mystique surrounding fire, its ambivalent nature, and 

its importance misled ancient Greeks and Chinese into 

identifying this powerful force as one of the four basic 

elements governing our planet; even today we are still 

trying to unravel the mysteries of this phenomenon. 

Luckily, complete knowledge is not a prerequisite to 

its effective use. Over time, our skill and ability to 

manipulate the behavior and effects of wildland fire has 

resulted in significant economic and societal gains.

This is not the place for an in-depth discussion of the 

economics of prescription fire because the overriding 

mandatory reason we must protect and facilitate its use 

is that it is an “Ecological Imperative.” That reason aside, 

prescription fire remains the least expensive method 

for modifying the landscape and as such always will be 

popular (e.g., see Dubois and others 2003). Alexander 

and Thomas (2006) described current costs of fuel 

treatment, including fire, while Wade and Moss (1999) 

showed that the periodic use of prescription fire results 

in a benefit/cost ratio of nearly 2 to 1—adding a return 

of more than $3 per acre per year for a landowner 

managing for southern pine. See Dale and others (2005) 

for a discussion of the need to factor the benefits of fire 

into wildfire suppression decisionmaking; they used 

a Colorado case study to show how substantial cost 

savings accrue from expanding the use of prescription 

fire. Mason and others (2006) provided an economic 

assessment that shows substantial net benefits from 

investments in fuel removals to reduce the incidence of 

crown fire. The literature contains numerous accounts 

from throughout the country of wildfires dropping to 

the ground, causing less damage, and becoming much 

easier to suppress after spreading into areas where 

fuelloads had been reduced by a previous fire (e.g., 

Moore and others 1955; Davis and Cooper 1963; 

Cumming 1964; Helms 1979; Wagle and Eakle 197;, 

Martin 1988; Ferguson 1998; Outcalt and Wade 2000).

When trying to measure the success of fire prevention, 

it is impossible to know how many fires your efforts 

prevented; the same holds true in determining what 

a wildfire would have cost to suppress and how much 

damage it would have caused had it not run into a 

relatively recent burn. Saveland (1987) used breakeven 

analysis to demonstrate that “large financial gains” 

would accrue from the use of prescription fire to reduce 

fuels in critical areas, and Jonathan Yoder is working on 

a dynamic economic model for prescription fire that 

incorporates risk, liability, and timing (e.g., Yoder 2004; 

Yoder and Blatner 2004).

LEARNING FROM THE PAST

We cannot look into the future to see the results of 

alternative actions before they are taken, but we can 

learn from the past. As former Florida State Forester 

John Bethea said: “You can no more get to where you 

don’t know where you’re going than you got to where you 

think you are from where you don’t know where you’ve 

been.” With that in mind we take a brief look at past fire 

management and legislative attempts to exclude fire.
2	 Fire 36 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

By the mid-19th century, steam, and early in the 20th 

century, the internal combustion engine, both with their 

tightly-harnessed fire, became commonplace and fire 

no longer was the only mechanism available to humans 

that could modify the environment on a landscape scale. 

Using these new tools, humankind changed the landscape 

at an awesome pace, but we could not distance ourselves 

from fire. We created an unprecedented accumulation 

of debris as we cleared the wilderness, clearcutting and 

high-grading our way across the continent. Nature’s 

match coupled with our own carelessness with fire, 

including the plethora of accidental fires resulting from 

these new inventions (especially steam locomotives, which 

spewed a never ending trail of sparks from their stacks), 

led to devastating fires. The result was conflagrations 

and tragedies like the Miramichi Fire that blackened 

more than 3 million acres of Maine and New Brunswick 

in 1825 with a loss of more than 160 lives; the 1871

Peshtigo Fire in Wisconsin, which killed 1,500 people 

and devastated 1,000 square miles of pine forest in 8 

hours; the 1903 Adirondack Fires; the postlogging fires of 

the Lake States in 1910 and 1918 which snuffed out 400 

lives; and the 1910 Idaho and Montana fires that burned 

3 million acres (USDA For. Serv. 1954).

Those fires led to a public outcry for change. The 

answer was obvious, or so it seemed; simply exclude 

the fire demon from wildlands and forests would at 

last be free to grow to their full potential, yielding an 

unprecedented bounty for human enjoyment and use. 

Although scientists, educators, and land managers 

recognized the need for suppressing unwanted fires, 

many also advocated the use of prescribed burning as a 

hazard reduction measure and fervently warned of the 

dangers of attempted fire exclusion. Gifford Pinchot 

(1898), the first Forest Service Chief and the person who 

implemented the Agency’s fire control policy, recognized 

the need for intentional fire to keep hazardous fuels 

from accumulating. But it was not to be; once the public 

spoke in a unified voice, Pinchot acquiesced even though 

he must have realized that the plan he implemented 

would not be successful over time. Abraham Lincoln put 

it, “Public sentiment is everything. With public sentiment, 

nothing can fail. Without it, nothing can succeed” 

(www.brainyquote.com/quotes/authors/a/abraham/

lincoln.html).

Thus began the federal fire exclusion policy and the 

federal government embraced it with zeal. Many forestry 

officials in the South recognized the folly of fire exclusion 

and did not actively endorse it. But despite this initial 

lack of support, the federal fire exclusion policy was 

extended to all lands with passage of the Clarke-McNary 

Act in 1924. This act used the carrot-and-stick approach 

(See Schiff 1962) that eventually forced all states to join 

the parade to a war that could not be won.

To make sure everyone is on the same page, fire 

suppression and fire entirely are completely different 

concepts. The objective of fire suppression is to 

extinguish unwanted/ illegal fires quickly in a safe, 

efficient manner; wildland fire suppression has been 

and doubtless will continue to be a high priority for all 

land management agencies; its long-term consequences 

are reduced damage to ecosystems, reduced financial 

losses to landowners, and increased public health and 

safety. Fire exclusion is the attempt to remove fire 

from the landscape; its long-term consequences are 

escalating suppression costs, decreased probability of 

success, increased risk to firefighters and the public, and 

unwanted ecosystem changes.

Most fire management agencies in the South believe that 

protecting the public and sustaining ecosystems entails 

a full-fledged fire prevention campaign, highly trained 

state-of-the-art fire suppression forces, and the extensive 

use of prescribed fire. Outside of the South, some states 

are slowly moving in this direction. For example, in 

2005, Michigan joined Florida, Georgia, and Nevada in 

providing landowners with the highest level of liability 

protection. The small steps many states are taking to 

include prescription fire as an agency mandate should be 

applauded rather than criticized for their slow progress. 

Such fundamental changes in philosophy should be 

incorporated operationally in well-planned small steps to 

build confidence and develop expertise.

The application of fire on the landscape is as much an 

art as a science because the behavior and effects of fire 

are microsite-specific and change as burning conditions 

change throughout the day and year. Many resource 

managers would rather not use fire because of the risks 

involved, but they do so because they recognize fire is 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 237

inevitable and that one is much better off using it under 

conditions they select as opposed to allowing nature and 

a temporally unpredictable ignition source to determine 

its timing. Bob Cooper, the first Project Leader at the 

Southern Forest Fire Laboratory, said “fire makes a good 

servant but a poor master.” Herbert Stoddard (1961), 

an early advocate of prescribed fire and the founder of 

Tall Timbers Research Station stated: “Fire may well be 

compared to a two-edged sword which requires judgment, 

care, and experience to properly handle....”

THE LEGACY OF FIRE EXCLUSION

On areas where fires had once been frequent, they were 

easy to extinguish due to a picturesque, low-stature, 

herbaceous ground cover. But as fuels accumulated on 

unburned areas and woody brush shaded out grasses and 

forbs, fires became more difficult to start, though once 

ignited they became increasingly difficult to suppress. 

When uncharacteristically high fuel accumulations 

inevitably did burn, rather than acting as a cleansing or 

rejuvenating force, fires often became high-intensity and 

catastrophic conflagrations. Systems that traditionally 

were perpetuated by fires spreading across the forest 

floor now faced fires that climbed into and consumed 

the canopy. Alterations in fire regime can lead to atypical 

successional pathways and extirpation of flora and fauna 

(Gill and Bradstock 1995; Glitzenstein and others 2003). 

Two early researchers, Frank Craighead during World 

War I in California and Harold Weaver several decades 

later in Oregon, documented the dramatic difference in 

forest health between forests burned on a regular basis 

where insects were endemic and those on surrounding 

unburned lands where insect populations were epidemic 

(Weaver 1959; Craighead 1977).

According to Wuerthner (1995): “No single human 

modification of the environment has had more pervasive 

and widespread negative consequences for the ecological 

integrity of North America than the suppression of fire. 

Fire suppression has destroyed the natural balance of the 

land more than overgrazing, logging, or the elimination 

of predators.” This opinion may represent one end of 

the spectrum but it should be obvious to people at the 

other end that the primary byproduct of attempted fire 

exclusion was the unprecedented and unnatural buildup 

of fuels on the Nation’s wildlands, which contributed 

to conflagrations capable of killing everything in their 

path. Vogl (1976) warned us that “Nature strives to 

maintain balances. Nonliving and living systems tend to 

be oriented toward a balanced state. When the existing 

quasi-equilibrium of these ecosystems and the organisms 

comprising them is disturbed or upset, feedback mechanisms 

come into play and phenomenal forces are amassed in the 

recovery and restoration to stability and balance.” This is 

exactly the scenario that unfolded with attempted fire 

exclusion in many sections of the country. Dead fuels 

accumulated and woody species previously restricted 

to the ground layer grew to form a dense midstory that 

shaded out the herbaceous ground cover and provided 

a pathway for fire to reach the canopy layer. From an 

ecosystem standpoint, the removal of fire jeopardized 

the long-term perpetuation of fire-adapted ecosystems. 

Frances Bacon understood the folly of the sort of 

thinking behind fire exclusion when he observed that 

“Nature is not governed except by obeying her” (http://

www.brainyquote.com/quotes/authors/f/francis_bacon.

html). Removing fire also violated principles described 

by Aldo Leopold (1949) in his A Sand County Almanac 

such that “The first rule of successful tinkering is not 

throwing away any of the pieces.” Fire exclusion amounted 

to throwing away one of the critical pieces.

Over time, the American public came to understand the 

unwelcome but inevitable consequences of attempted fire 

exclusion and realized that we do not have the ability to 

stop catastrophic fires once they start. When faced with 

such fires, firefighters have to wait for a change in the 

weather, a change in topography, or for the fire to run 

out of fuel. The only practical alternative is to alter the 

fuels that foster these fires.

Although it became more obvious with each passing 

decade that the marriage between fire and wildlands 

could not be split asunder, it took 70 years before the 

public finally demanded a stop to government efforts 

to divorce them. Despite the outcry, which resulted in a 

new federal policy that now includes the use of fire, e.g.,

U.S. Environ. Prot. Agency (1998), U.S. Dep. Interior 

and others (2001), USDA For. Serv. (2003), the debate 

continues as we cope with catastrophic fires and the 

legacy of this failed exclusion policy, and ponder how 

enthusiastically we should embrace and regulate the 
2	 Fire 38 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

intentional use of wildland fire. Several federal incentive 

programs promote and facilitate its use, for example, the 

Healthy Forest Restoration Act, Landowner Incentives 

Program and Wildlife Habitat Incentives Program. 

But this encouragement to burn lacks the same zeal 

or determination once used to promote fire exclusion. 

Also, these programs often seem to place more emphasis 

on fuel reduction than on the restoration of a natural 

process. See McCarthy (2002a) for a discussion of the 

politics involved.

Ten years after this watershed change in fire policy, 

it seems that there has been considerable vocal but 

relatively little operational progress, even in the South. 

The 13 Southern States contain about 225.6 million 

acres of forest land, 219 million of which are available 

for commercial use. This commercial forest land 

comprises 70 million acres of pine, 32 million acres 

of mixed pine/hardwood, and 117 million acres in 

hardwood. About 11 percent or 24.8 million acres of 

this is government owned (Gramley 2005). We estimate 

at least an additional 100 million acres, most of which 

is in Oklahoma and Texas, is in unimproved and 

improved rangeland/pasture. If Frost’s (1998) estimate of 

presettlement fire regimes is correct, the enormity of the 

task before us becomes clear. According to Palmer and 

others (2004), only about 35 percent of state lands and 

1 to 5 percent of private forestlands are being burned as 

often as they need to be for proper fuels management.

Few if any managers have the staff, funding, or full 

commitment of their superiors necessary to treat all the 

acres needing fire given the limited number of days with 

acceptable weather and fuel conditions. One impediment 

is the lack of incentives for a fire manager to take risks. 

It is much easier and professionally safer to advocate the 

benefits of fire but to never have the “right” conditions 

than it is to shoulder the risks involved with authorizing 

or conducting a burn. Prescribed fire requires action and 

with action comes responsibility. Fire exclusion is often a 

do-nothing-then-react approach that ensures anonymity 

and protection from the responsibility of taking action. 

There is always more responsibility associated with 

taking action than with reacting to outside stimuli. 

Although the risk of a bad outcome decreases with 

multiple burns on an area, it never disappears and the 

law of averages always is at work. Currently, there are few 

if any incentives for “doing the right thing” other than 

one’s own belief that as a manager “it is my obligation 

and moral responsibility to future generations to try to 

maintain healthy, fully functional ecosystems.” In the 

administrative climate of most organizations, including 

every federal land management agency, the penalties for 

a bad outcome let alone a mistake can be so severe that 

the only prudent choice for some professionals is not to 

burn. When fire is unleashed carelessly or with harmful 

intent, it can have devastating consequences. Thus, 

prescription fire carries with it a huge responsibility 

to use it wisely. Fire managers willing to shoulder this 

responsibility deserve the protection a statute can offer.

Nationally, we are losing fire experience to retirements 

faster than it can be replaced. This loss of experience 

places the entire prescribed fire industry at greater risk 

because burners with less experience are called upon to 

burn in a world far more complex than the one in which 

the “old fire dogs” gained invaluable experience. We 

cannot afford to exacerbate this problem by discouraging 

fire managers because the risks become too high for them 

to bear absent a good prescribed fire law.

Prescribed fire also needs to be protected from those who 

want to eliminate it for nonscientific personal reasons 

ranging from air quality issues to a misguided belief 

that fire is patently bad for plants and animals. If people 

opposed to the intentional use of fire become organized 

in the absence of a good law promoting/protecting 

prescribed fire, this valuable tool can be lost. That came 

close to happening in Georgia in 1989 when an escaped 

agricultural burn ignited peat adjacent to Interstate 75 

and “smoked in” the I-75 for several weeks while the 

state legislature was in session. The governor demanded 

action and a bill was quickly introduced to ban all open 

burning. Although it was defeated, prescribed burners 

in Georgia got the message and a bill modeled after the 

Florida Act was introduced and passed the following 

year. It occurred again in Georgia 10 years later when 

the Environmental Protection Agency (EPA) threatened 

to withhold federal highway funding if Georgia failed 

to develop a strategy to reduce air pollution in Atlanta 

resulting primarily from the 100 million miles driven 

per day in the area (Leinberger 1998); Pollard 2003; 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 239

Fed. Highway Admin. (2003). This megalopolis is the 

fastest growing human settlement in history and by 

most measures the most sprawling major metro area in 

America because Atlanta has chosen to grow out rather 

than up. Rather than curtailing this metastasis, Georgia 

demonstrated its unwillingness to make the hard choice 

and instead restricted open burning in the surrounding 

43 counties from May through September. This policy 

is severely hampering efforts to restore the rare montane 

longleaf pine ecosystems of northwest Georgia, which 

require growing-season burns.

The problem with such lack of moral fortitude is 

that it is contagious. For example, in 2004, the EPA 

added Augusta, Columbus, and Macon to its list of 

nonattainment areas in Georgia. Many people were 

blindsided by the Georgia Environmental Protection 

Division (GAEPD), which unveiled its plan to appease 

the EPA by using the same strategy used in Atlanta, 

namely, the elimination of all open burning during the 

months of May through September in an 11-county 

area, which includes a national wildlife refuge, a national 

forest, and two large military bases, all of which depend 

on growing-season prescription fires for hazard reduction 

and T&E species management. Agricultural burns 

were to be exempted and when GAEPD was asked to 

explain the reason for the agriculture exemption, officils 

stated: “They have a more powerful lobby.” Further, 

GAEPD considers the contribution of carbon dioxide 

from agricultural burning as irrelevant because “the 

carbon dioxide released is reabsorbed by crop regrowth in 

the next growing season” (Georgia Dep. Nat. Resour. 

1999). Using published GAEPD estimates that showed 

combustion of biomass contributes less than 1/10 of 1

percent of greenhouse emissions in Georgia, compared 

to 48 percent from petroleum and 44 percent from coal 

(there are two coal-fired power plants in the Macon 

vicinity) and that forests sequester more carbon dioxide 

each year than is produced by all sources of biomass 

combustion, (Georgia Dep. Nat. Resour. 1999), these 

proposed restrictions were scrapped and a compromise 

reached. Because such horror stories are all too common, 

we must be continually on guard and prepared to 

intelligently refute such tripe within the short time frame 

constitutionally allowed before it takes effect. Part of this 

preparation is to have a prescribed fire statute that makes 

prescription fire a landowner right and includes language 

to shield landowners from corrupt and/or misguided 

officials. Fire councils can demonstrate their usefulness in 

such situations by providing a prompt, recognizable, and 

unified voice for prescribed fire.

WHY WE NEED LAWS 

PROTECTING AND FACILITATING 

PRESCRIPTION FIRE

o	The general public, to a large degree, suffers 

from the misconception that “all fire is bad” 

and thus views the intentional use of fire as 

counterproductive at best.

o	Prescribed fire is a resource management tool 

that benefits the safety of landowners and the 

public, the environment, and the economy of 

community, state, and nation.

o	There are risks associated with the use of fire:

	Escapes that can damage another’s property.

	Smoke intrusions that can cause health and 

safety concerns.

	These risks can be due to negligence on the 

part of the burner or due to unpredictable 

events.

	Burners should be held responsible for 

damages when they do not follow appropriate 

standards, particularly where spelled out in 

the law.

o	Burners should have personal protection from 

damages to others when appropriate precautions 

were taken and due caution exercised.

o	Burners should be protected from nuisance 

complaints when they follow appropriate burn 

execution standards.

o	Burners should be rewarded for enhancing their 

skills and ability to excel in their use of this risky 

undertaking.

o	Without such a law, some resource managers 

“talked the talk” but continually found excuses 

not to burn. When nature does it, they are not 

held accountable.

Good rules to facilitate fire use will make the planning 

and execution of a burn more efficient and cost effective 
2	 Fire 40 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

by eliminating needless and often repetitious paperwork 

requirements and multiple approvals, by recognizing 

and rewarding practitioners who strive to increase their 

knowledge and skills, and by creating a favorable legal 

environment that provides meaningful burner protection.

Neil Sampson (1995), executive vice president of The 

American Forestry Association, stated: “Fire is the most 

powerful, unpredictable, and potentially deadly tool land 

managers can use.” A point of clarification is that fire 

behavior is predictable but the predictions are only as 

accurate as the inputs, for example, the weather forecast 

they were based on. See Weiner (1985), Siegel (1986), 

and Stanton (1995) for a discussion of the legal aspects 

of prescribed burning. We believe resource managers 

should be held more accountable for not using fire in 

fire-adapted ecosystems than for using this “Ecological 

Imperative” and experiencing a bad outcome that is 

beyond their control.

Because of the ecological necessity of periodic fire, the 

cultural heritage of managed fire, and the threat to fire 

use resulting from an increasing population, southern 

states have sought legislation to protect the future of 

prescribed fire as a management tool, but they are not 

alone. Sun (in press) reviewed the liability burdens placed 

on landowners who use prescription fire in all 50 States 

and recent statute changes. He found that all but six state 

statutes have evolved from “heavy liability burdens” on 

landowners using prescription fire toward a negligence 

tort rule (Appendix C), probably due to a resurgence in 

demand for prescription fire as a management tool. Easyto-understand descriptions of the differences between 

strict tort liability, uncertain liability, simple negligence, 

and gross negligence are also given. Currently, 22 states 

have legislation protecting/encouraging the use of 

prescribed fire. Seventeen are east of the Great Plains, 

including 11 in the South: Alabama, Arkansas, Florida, 

Georgia, Louisiana, Mississippi, North Carolina, 

Oklahoma, South Carolina, Texas, and Virginia.

THE HUMAN FACTOR

Given the obvious need to drastically increase the 

amountof acreage treated with fire, what is the holdup? 

The simple answer is that wildland fire and people do 

not mix well and the South has the fastest growing 

population in the United States. But as Stowe (2004a) 

pointed out, “While it is tempting to blame the increasing 

hassles associated with land management practices on these 

immigrants, we would do well to remember that many of 

these folks come from a region with a recent fire history 

much different than that of the South.” Immigrants from 

the North are joined by affluent southern urbanites who 

want to escape the pollution and closeness of city life for 

a home in the woods surrounded by nature. Matthews 

(1992) defined the term “rurbanization” as “the invasion 

of affluent urban and suburban-oriented people into rural 

areas, looking for a self-defined ‘country’ lifestyle, while 

importing urban attitudes and values and expecting urban 

amenities.” These rurbanites generally are unaware of 

the hazardous buildup of fuels in their own backyard 

(Gardner and others 1987). Most view just blackened 

landscapes as damaged rather than as rejuvenated. 

Southern states have multifaceted outreach programs to 

educate new arrivals and their children on the biological 

necessity of wildland fire in the fuel types where they 

have chosen to live. But when asked about his agency’s 

outreach regarding prescribed fire, Florida State Forester 

Mike Long lamented: “They are immigrating faster then 

we can educate them.”

Many rurbanites will tell you that they desire intact, fully 

functional ecosystems, including the judicious use of 

prescribed fire, but in reality their acceptance often is on 

a conceptual rather than on an operational level. Most 

do not like the smell of smoke, being inconvenienced by 

ashes in their swimming pools, or subjected to detours 

and delays when traveling because of smoke on the road. 

This situation will continue to deteriorate so long as 

people choose to build out rather than up. Many people 

have a strong desire to live in or at the edge of wildlands. 

But this places people and their homes among fire-prone 

vegetation. People living at this wildland urban interface 

(WUI) do not realize they can, or believe they should 

have to, reduce fuels around their homes to make them 

more fire safe; they expect the taxes they pay to provide 

complete protection even when unnatural, fuel-infested 

plant communities catch fire. The truth is that when fire 

enters the WUI, suppression forces have to make hard 

choices regarding the allocation of resources. Firefighters 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 241

must allocate resources to evacuate residents and protect 

unprepared houses, both at the expense of resources 

that could have been used to combat the spread of the 

fire. When resources for suppressing fire are curtailed, 

it grows larger and threatens even more homes. People 

who choose to live in the WUI must realize that fire 

protection is not a certainty and that they must shoulder 

their share of the responsibility for protecting their 

homes.

Insurance rate structures are used to encourage better 

driving habits and to encourage homeowners to protect 

their homes from theft. Similar rate structures could 

be used to encourage homeowners to shoulder more 

of the responsibility to reduce the threat of wildfire on 

their property. If the insurance industry was forced to 

determine rates based on risk in a particular area rather 

than spreading the cost of insuring homes in high firerisk areas across its entire domain, and if homeowners 

using wood shakes for roofing or siding or plastic soffit 

vent covers were unable to collect in the event the 

structure was damaged by wildland fire, we believe that 

problem would solve itself quickly. Another problem 

is that disaster loans and low premiums subsidize 

inappropriate and high-risk construction (Davis 1990).

State fire management agencies are committed to work 

with the public to make the interface a safer place to 

live, but they cannot do it alone. As fires in the WUI 

became more problematic, this issue has been addressed 

through land use planning, zoning, structural codes, and 

development design (Irwin 1987; Davis 1990; Rice and 

Davis 1991; Pumphrey 1993; Haines and Cleaves 1995). 

But as Monroe (2002) lamented: “Unfortunately, few of 

these recommendations are easy to implement after homes 

are built, and few seem to be heeded during the planning 

phase.” On a positive note, this affords an opportunity 

for local and state governments to help safeguard 

their constituencies by providing guidance to and 

requirements for developers, builders, and homeowners. 

State and local officials should be encouraged to seek 

federal funding support for fuels mitigation and for the 

use of federal prescribed fire use teams on nonfederal 

public and private lands. See Plevel (1997), Monroe 

(2002) and Long and others (2005) for suggestions.

IDENTIFYING THE RIGHT PEOPLE TO 

FACILITATE PRO FIRE LEGISLATION

Once the need for a prescribed fire law is recognized and 

the decision to craft a bill is made, the next step is to 

get the right people involved. To enhance the chances of 

success, you would do well to marshal a diverse group 

of fire practitioners that includes state wildfire and 

wildlife agencies; all state and federal land management 

agencies; the structural fire community; community 

planners; the insurance industry; municipal and county 

land managers; forest industry; the state forestry 

association; range management groups; nongovernment 

organizations involved in land management and 

restoration such as The Nature Conservancy; private 

landowners who use fire; and sportsman’s clubs and 

other special interest groups such as the National Wild 

Turkey Federation. By creating a united front, the team 

can demonstrate this is a larger issue than any one entity. 

The team can also send the right messenger, or group 

of messengers, to meet with people who oppose the 

proposed legislation. A team also can deflect attacks on 

an individual member. Southern cattlemen and farmers 

are major contributors to the acreage burned with 

intentional fire, but both groups have initially tended 

to choose not to be covered under prescribed burn 

legislation because it would be much more restrictive to 

them. However, the situation may differ in your state, 

so we suggest you approach such groups because they 

generally have strong lobbies.

Every restriction to prescribed burning means fewer 

acres treated. The underlying purpose of a prescribed fire 

law is to help ensure that the assemblages of indigenous 

plants and animals that make up our wildlands continue 

to thrive as fully functional ecosystems. Legislation 

cannot make it so but it can and should promote, 

facilitate, and protect the intentional use of fire for 

the benefit of both private landowners and society as a 

whole. Current laws and attendant rules and regulations 

in many states severely curtail the use of prescription 

fire; in some states, open burning is banned during the 

day while in others the permitting process is a nightmare 

(see McCarthy and Foster 2002). When drafting pro-fire 

legislation, include language that addresses contradictory 

language in existing law.
2	 Fire 42 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Prescribed fire legislation should be drafted so that 

it both facilitates fire use and allows for adaptive 

management. We list a number of items/issues that 

should be considered when drafting pro-fire legislation 

(Appendix F). We do not want to once again enact fire 

statutes that dictate a specific course of action only to 

find later that they were flawed and/or shortsighted. 

An excellent strategy to assure this does not happen is 

to make such legislation general and have the statute 

delegate authority to a state agency to formulate the 

details through its rule-making process (See Appendix B). 

Another reason to keep the law general is that support 

for a broad concept can be developed by reaching out to 

a diverse group of organizations, but persuading these 

groups to reach a consensus on the details of such a law 

is difficult at best. Such a strategy enables the agency 

to practice adaptive management and make changes 

as warranted without exposing the statute itself to the 

fickle nature of legislatures and the political necessities 

of election year politics. It took nearly a century of fire 

exclusion to create the current situation and it will take 

decades to resolve the problem. Conditions will continue 

to change and new tools will become available.

It is essential that we take action because the no-action 

approach will not resolve this issue despite what some 

people want to believe. Vitousek and others (1997) 

described the necessity of active management well when 

they wrote: “There is no clearer indication of the extent of 

human domination of Earth than the fact that maintaining 

the diversity of ‘wild’ species and the functioning of ‘wild’ 

ecosystems will require increasing human involvement.”

Howard (1974) believed it is our obligation: “Man has a 

moral responsibility to manage nature once he disrupts it.”

That belief was echoed by Dr. Patrick Moore, cofounder 

and former president of Greenpeace, who stated: “...we 

have a responsibility/obligation to use our knowledge and 

experience to keep US forests healthy.”

There will be people who resist any action taken to 

manage natural lands. Leopold (1942) identified the 

root of the problem that causes well-meaning people 

to fight the very activities needed to perpetuate fully 

functional ecosystems. He wrote: “I am convinced that 

most Americans have no idea what a decent forest looks 

like. The only way to tell them is to show them.” There are 

numerous examples of communities where succession 

has been stabilized at a particular sere by chronic lowintensity fires, as well as examples of those that have 

been rejuvenated by having succession set back to an 

earlier successional stage with less frequent but higher 

intensity fires. The benefits of the proper application of 

fire can be seen on these sites which serve as standards 

we can emulate. But without the continued presence 

of fire they too will disappear. If we do not begin to 

use fire on a much grander scale, there will be far fewer 

of these sites to behold. It would be a shame if we end 

up with nothing worthwhile to show (Orr 1993). You 

can contact any of the authors for information on the 

location of healthy, fire maintained communities.

Consider organizing a fire council prior to trying 

to enact prescribed fire legislation. Fire councils are 

excellent forums for disseminating knowledge, and an 

excellent way to keep prescribed burners current on 

new research results, emerging technology, training 

opportunities and other fire-related issues in your state 

and beyond. A number of southern states have a fire 

council (Florida has three). If there is no fire council in 

your area, we urge you to form one (see Miller 1998).

PUBLIC OPINION CAN LEAD TO 

PRO-FIRE LEGISLATION

The public is confused because it continues to receive 

mixed messages. On the one hand, resource professionals 

are trying to educate the public about the benefits of 

the judicious use of fire and its biological necessity. On 

the other hand, newspapers and television often focus 

on destructive fires resulting from years of attempted 

fire exclusion. The dilemma that faced the Smoky Bear 

advertising campaign perhaps exemplifies the situation. 

It’s been said this was the most successful ad campaign 

in history; since 1944 Smokey has convinced three 

generations of Americans that all forest fires are patently 

bad. On Smokey’s website (www.smokeybear.com/), you 

now can find a link to a discussion of the intentional 

use of fire, though it is not well identified. In fact, the 

word “fire” is removed from the title. Smokey has never 

said he was wrong or apologized for misleading the 

public, and his website includes nothing that suggests 

he misinformed us, so it’s no wonder much of the 

public is still confused. He needs to take a cue from 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 243

his colleague Mark Trail and fully embrace prescription 

fire. One reason we are so adamant about Smokey is 

because it is still commonplace to see his suggestio falsi on 

posters and billboards. It is these falsehoods that create 

perceptions and opinions. According to the Wildland [fire] 

Communicator’s Guide (National Interagency Fire Center, 

http://www.nifc.gov/preved/comm_guide/wildfire/index.

html), which is linked to Smokey’s website: “...these 

perceptions, often misperceptions, can become inculcated 

into society to form ‘public opinion’. That public opinion 

influences laws and policies related to wildland fire 

management.” It is up to all of us to try to educate the 

public and replace Smokey’s flawed message with the 

truth. But this is not an easy task; for example, two of 

the coauthors were threatened with jail for putting a 

driptorch in Smokey’s hand a number of years ago.

Our primary message is to follow President Lincoln’s 

advice and educate the public so they also can reach the 

same conclusion as fire professionals, i.e., that wildland 

fire is truly the “Ecological Imperative.” You will bolster 

your chances of success by using published scientific 

evidence and existing pro-fire legislation from other 

areas to support your case. If you are successful in these 

outreach efforts, your ultimate goal of getting pro-fire 

legislation passed will be much easier.

We also need to look inward at our everyday activities 

to see what we can do to promote fire use. Redford 

and Tabor (2000) observed: “Conservation practitioners 

rarely write about the work they do,” especially their 

failures. “Renewal of funding is contingent upon success. 

Few have ever been rewarded for anything other than 

success. Inside this straightjacket we will not achieve 

effective conservation because we will never learn. Learning 

requires experimentation and experimentation sometimes 

means failure. When failure is not tolerated learning will 

never take place. This situation in which experimentation, 

failure, and learning are not tolerated is a death spiral 

for conservation. We and all we are trying to save will 

not survive if we do not break out of this inward-turning 

spiral and move into the uneven and unpredictable terrain 

of a highly self-critical adaptive management approach. 

Time is short as we try to slow the juggernaut of biotic 

impoverishment. We cannot waste time trying things that 

others have tried and found wanting. But we cannot do 

otherwise unless we all document our failures as well as our 

successes. What is needed is a ‘safe-fail’ environment where 

folks are encouraged to innovate, experiment, and learn; 

but most importantly to document what has been tried, 

and what has failed. We suggest that the long term success 

of conservation depends on willingness not only to admit 

our failures but to share them as well.” As practitioners, 

we need to take the time to record what we have found 

that works and what does not, and disseminate that 

information.

Most bureaucrats and elected officials at all levels of 

government will publicly state that they are against the 

extinction of ecosystems. Given that fire is the Ecological 

Imperative necessary for the survival of fire-adapted 

ecosystems, it is in the best interest of public officials 

to demonstrate their support for the judicious use of 

prescribed fire by protecting it as a landowner right 

and by facilitating its use. South Carolina passed the 

Heritage Trust Act in 1976 (www.scstatehouse.net/code/

t51c017.doc), the first law of its kind that specifically 

provided for prescription fire. Florida passed a law in 

1977 that required absentee landowners to reduce fuel 

accumulations judged to be hazardous, or reimburse the 

state for doing so (Wade and Long 1979). The provisions 

of this statute were embodied in the 1999 changes 

(Brenner and Wade 2003) to the 1990 Florida Prescribed 

Fire Act (Brenner and Wade 1992) (Appendix A) 

wherein Florida followed the example set by Nevada in 

1993 and raised its liability standard to gross negligence.

Walt Thomson with The Nature Conservancy in Florida 

pointed out a shortcoming in the Florida Prescribed Fire 

Act in that it offers safe harbor to prescribed burners 

through the standard of care level for gross negligence 

This essentially dictates that if you are trained and 

become certified, you are allowed to exercise a lower 

standard of care. He suggests that “safe harbor” should be 

gained through other strategies, e.g., levels of exemption 

or liability caps. He raises a valid point but one that 

we believe can be handled by the Florida Fire Control 

Bureau through its oversight of certified burners. For 

example, it could require certified burners to report 

all fire escapes that leave the landowner’s property or 

that result in a smoke incident to the Bureau, which 

would then use established procedures to review the 
2	 Fire 44 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

situation and strip the burner of his or her certification 

if warranted, thereby removing that individual’s gross 

negligence protection.

At the local level, all 67 counties in Florida have passed 

resolutions supporting prescription fire (Appendix D) 

and 48 counties have ordinances stating that the use of 

prescribed fire is a property owner’s right. Some counties, 

e.g., Flagler have ordinances that require property owners 

to clear “nuisance brush and pine” trees or else reimburse 

the Division of Forestry for the service; others, such 

as Sarasota County, have created smoke corridors for 

prescription burns and require realtors to provide a copy 

of such to all perspective property buyers within these 

corridors (Appendix E).

More in-depth discussions of Florida laws pertaining 

to prescription fire and accompanying administrative 

rules are found in Brenner and Wade (1992), Wade 

and Brenner (1995), and Brenner and Wade (2003). 

Both the Florida and Georgia prescribed burn laws have 

withstood challenges.

It should be noted that there are encouraging signs 

regarding wildland fire in other regions of the Nation, 

especially in educating and enticing landowners to take 

more responsibility in protecting their property from 

fire. McCarthy (2002b) described an innovative effort to 

educate homeowners undertaken by the Lands Council 

in Spokane, Washington. The nationwide FireWise 

program (www.firewise.org) and Canadian Partners in 

Protection (www.partnersinprotection.ab.ca/) are others. 

CONCLUSION

Ultimately it will be the public that decides whether 

intentional fire will be used to manage fire-adapted 

ecosystems. We all must be proactive in outreach 

efforts to demonstrate that the public and our elected 

representatives that prescribed fire is the only practical 

(and rational) approach, and that given the latitude and 

legal protection to do so, fire managers have the will and 

skill to use fire in a safe and effective manner. It is now 

up to all of us to rise to the challenges ahead and to take 

action rather than hide from the challenge and risks and 

be reduced to simply reacting. The cost of no action is 

simply too great.

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Wade, D.; Moss, S. 1999. The economics of 

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Manuscript on file West Virginia University.

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List of Appendices:

A. Florida Prescribed Fire Act

B. Florida Division of Forestry Rules and Regulations 

pertaining to fire 

C. Prescribed fire liability rules retained on U.S. forest 

land by state 

D. Sarasota County, Florida resolution supporting 

prescribed fire

E. Sarasota County, Florida Notice of Proximity 

establishing smoke corridors

F. Items to consider when drafting a prescribed fire 

statute
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Appendix A: The Florida Prescribed Burning Act. A model state 

statute to protect and promote the use of prescription fire.

590.125 Open burng authorized by the division.—

(1) DEFINITIONS.—As used in this section, the term: 

(a) “Prescribed burning” means the controlled application of fire in accordance with a written 

prescription for vegetative fuels under specified environmental conditions while following appropriate 

precautionary measures that ensure that the fire is confined to a predetermined area to accomplish the 

planned fire or land-management objectives.

(b) “Certified prescribed burn manager” means an individual who successfully completes the 

certification program of the division and possesses a valid certification number.

(c) “Prescription” means a written plan establishing the criteria necessary for starting, controlling, and 

extinguishing a prescribed burn.

(d) “Extinguished” means that no spreading flame for wild land burning or certified prescribed burning, 

and no visible flame, smoke, or emissions for vegetative land-clearing debris burning, exist.

(2) NONCERTIFIED BURNING.

(a) Persons may be authorized to burn wild land or vegetative land-clearing debris in accordance with 

this subsection if: 

1. There is specific consent of the landowner or his or her designee;

2. Authorization has been obtained from the division or its designated agent before starting the 

burn; 

3. There are adequate firebreaks at the burn site and sufficient personnel and firefighting 

equipment for the control of the fire; 

4. The fire remains within the boundary of the authorized area; 

5. Someone is present at the burn site until the fire is extinguished; 

6. The division does not cancel the authorization; and 

7. The division determines that air quality and fire danger are favorable for safe burning. 

(b) A person who burns wild land or vegetative land-clearing debris in a manner that violates any 

requirement of this subsection commits a misdemeanor of the second degree, punishable as provided in 

s. 775.082 or s. 775.083.

(3) CERTIFIED PRESCRIBED BURNING; LEGISLATIVE FINDINGS AND PURPOSE.

(a) The application of prescribed burning is a land management tool that benefits the safety of the 

public, the environment, and the economy of the state. The Legislature finds that: 

1. Prescribed burning reduces vegetative fuels within wild land areas. Reduction of the fuel load 

reduces the risk and severity of wildfire, thereby reducing the threat of loss of life and property, 

particularly in urban areas.

2. Most of Florida’s natural communities require periodic fire for maintenance of their ecological 

integrity. Prescribed burning is essential to the perpetuation, restoration, and management of 

many plant and animal communities. Significant loss of the state’s biological diversity will occur 

if fire is excluded from fire-dependent systems.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 251

3. Forestland and rangeland constitute significant economic, biological, and aesthetic resources of 

statewide importance. Prescribed burning on forestland prepares sites for reforestation, removes 

undesirable competing vegetation, expedites nutrient cycling, and controls or eliminates certain 

forest pathogens. On rangeland, prescribed burning improves the quality and quantity of 

herbaceous vegetation necessary for livestock production. 

4. The state purchased hundreds of thousands of acres of land for parks, preserves, wildlife 

management areas, forests, and other public purposes. The use of prescribed burning for 

management of public lands is essential to maintain the specific resource values for which these 

lands were acquired. 

5. A public education program is necessary to make citizens and visitors aware of the public safety, 

resource, and economic benefits of prescribed burning. 

6. Proper training in the use of prescribed burning is necessary to ensure maximum benefits and 

protection for the public. 

7. As Florida’s population continues to grow, pressures from liability issues and nuisance 

complaints inhibit the use of prescribed burning. Therefore, the division is urged to maximize 

the opportunities for prescribed burning conducted during its daytime and nighttime 

authorization process.

(b) Certified prescribed burning pertains only to broadcast burning. It must be conducted in accordance 

with this subsection and: 

1. May be accomplished only when a certified prescribed burn manager is present on site with a 

copy of the prescription from ignition of the burn to its completion. 

2. Requires that a written prescription be prepared before receiving authorization to burn from the 

division. 

3. Requires that the specific consent of the landowner or his or her designee be obtained before 

requesting an authorization. 

4 Requires that an authorization to burn be obtained from the division before igniting the burn. 

5. Requires that there be adequate firebreaks at the burn site and sufficient personnel and 

firefighting equipment for the control of the fire. 

6. Is considered to be in the public interest and does not constitute a public or private nuisance 

when conducted under applicable state air pollution statutes and rules. 

7. Is considered to be a property right of the property owner if vegetative fuels are burned as 

required in this subsection. 

(c) Neither a property owner nor his or her agent is liable pursuant to s. 590.13 for damage or injury 

caused by the fire or resulting smoke or considered to be in violation of subsection (2) for burns 

conducted in accordance with this subsection unless gross negligence is proven. 

(d) Any certified burner who violates this section commits a misdemeanor of the second degree, 

punishable as provided in s. 775.082 or s. 775.083.

(e) The division shall adopt rules for the use of prescribed burning and for certifying and decertifying 

certified prescribed burn managers based on their past experience, training, and record of compliance 

with this section.
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(4) WILDFIRE HAZARD REDUCTION TREATMENT BY THE DIVISION.—The division may 

conduct fuel reduction initiatives, including, but not limited to, burning and mechanical and chemical 

treatment, on any area of wild land within the state which is reasonably determined to be in danger of 

wildfire in accordance with the following procedures:

(a) Describe the areas that will receive fuels treatment to the affected local governmental entity.

(b) Publish a treatment notice, including a description of the area to be treated, in a conspicuous 

manner in at least one newspaper of general circulation in the area of the treatment not less than 10 days 

before the treatment. 

(c) Prepare, and the county tax collector shall include with the annual tax statement, a notice to be sent 

to all landowners in each township designated by the division as a wildfire hazard area. The notice must 

describe particularly the area to be treated and the tentative date or dates of the treatment and must list 

the reasons for and the expected benefits from the wildfire hazard reduction. 

(d) Consider any landowner objections to the fuels treatment of his or her property. The landowner may 

apply to the director of the division for a review of alternative methods of fuel reduction on the property. 

If the director or his or her designee does not resolve the landowner objection, the director shall convene 

a panel made up of the local forestry unit manager, the fire chief of the jurisdiction, and the affected 

county or city manager, or any of their designees. If the panel’s recommendation is not acceptable to the 

landowner, the landowner may request further consideration by the Commissioner of Agriculture or his 

or her designee and shall thereafter be entitled to an administrative hearing pursuant to the provisions of 

chapter 120.

(5) DUTIES OF AGENCIES.—The Department of Education shall incorporate, where feasible and 

appropriate, the issues of fuels treatment, including prescribed burning, into its educational materials.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 253

Appendix B: FL Division of Forestry rules and regulations

pertaining to Appendix A

5I-2.006 Open Burning Allowed.

(1) Open Burning in General. Authorization must be obtained from the Florida Division of forestry 

for burns relating to agriculture, silviculture and pile burning on the same day the burn is to take 

place or after 4:00 p.m. of the previous day. The Division of Forestry will set special requirements for 

authorizations in order to protect public health and safety, including; on site inspections, restricting 

wind direction, limiting the burning period, halt or limit burning when fire danger is too high in 

all, or specific parts of the state, and requiring specific personnel and control equipment on site. Any 

authorized burn that goes out of compliance, but has not escaped the authorized area will be allowed a 

maximum of two hours to be brought into compliance by the person responsible. In the event that the 

Division determines that there is a threat to life, public safety or property, immediate suppression action 

will be taken by the Division of Forestry.

(a) Daytime Non-Certified Authorizations will be issued for the burning to be conducted from 8:00 

a.m. CT or 9:00 a.m. ET and the fire must discontinue spreading one hour before sunset.

(b) Nighttime Non-Certified Authorizations will be issued with a Dispersion Index of 8 or above for 

the burning to be conducted between one hour before sunset and 8:00 a.m. CT or 9:00 a.m. ET the 

following morning. Ignition of these fires is authorized up to midnight CT or ET (specific to the time 

zone where the fire is located), however the fire can continue to spread until 8:00 am CT or 9:00 a.m. 

ET the following day. If additional time is required a new daytime authorization must be obtained from 

the Division.

(2) Open Burning for Certified Prescribed Burn Managers (CPBM). (All burning conducted under 

this section is related to broadcast burning for the purposes of; Silviculture, Wildlife Management, 

Ecological Maintenance and Restoration, Range and Pasture Management. Open burning authorizations 

under this section require the Certified Prescribed Burn Manager’s certification number be presented at 

the time of the request, and that a Certified Prescribed Burn Manager be on site for the entire burn.

(a) Prescription. A prescription for the burn must be completed prior to any ignition and it must be 

on site and available for inspection by a Department representative. The prescription will contain, as a 

minimum, (unless agreed to in writing locally between the burner and the District or Center Manager of 

the Division of Forestry) the following:

1. Stand or Site Description;

2. Map of the area to be burned;

3. Number of personnel and equipment types to be used on the prescribed burn;

4. Desired weather factors, including but not limited to surface wind speed and direction, 

transport wind speed and direction, minimum mixing height, minimum relative humidity, 

maximum temperature, and the minimum fine fuel moisture;

5. Desired fire behavior factors, such as type of burn technique, flame length, and rate of spread;

6.The time and date the prescription was prepared;

7. The authorization date and the time period of the authorization;
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8. An evaluation and approval of the anticipated impact of the proposed burn on related smoke 

sensitive areas;

9. The signature and number of the Certified Prescribed Burn Manager.

(b) Open Burning Hours

1. Daytime CPBM Authorizations will be issued for the burning to be conducted from 8:00 a.m. 

CT or 9:00 a.m. ET and the fire must discontinue spreading one hour after sunset.

2. Nighttime CPBM Authorizations will be issued with a Dispersion Index of 6 or above for the 

burning to be conducted between one hour before sunset and 8:00 a.m. CT and 9:00 a.m. 

ET the following day. Ignition of these fires is authorized up to midnight, however the fire can 

continue to spread until 8:00 a.m. CT and 9:00 a.m. ET the following day. If additional time is 

required a new authorization (daytime) must be obtained from the Division. The Division will 

issue authorizations at other times, in designated areas, when the Division has determined that 

atmospheric conditions in the vicinity of the burn will allow good dispersement of emissions, 

and the resulting smoke from the burn will not adversely impact smoke sensitive areas, e.g., 

highways, hospitals and airports.

(c) Burn Manager Certification Process. Certification to become a Certified Prescribed Burn Manager is 

accomplished by:

1. Satisfactory completion of the Division of Forestry’s Prescribed Fire Correspondence Course and 

direct experience in three prescribed burns prior to taking the course or;

2. Satisfactory completion of the Division of Forestry’s Prescribed Fire Classroom version of the 

Correspondence Course and a minimum of managing three prescribed burns prior to taking the 

course or;

3. Satisfactory completion of the Florida Inter-Agency Basic Prescribed Fire Course and direct 

experience in three prescribed burns following successful completion of the classroom training. 

The burns conducted during the training do not count as part of this three burn requirement.

4. Applicants must submit a completed prescription for a proposed certifying burn to their local 

Florida Division of Forestry office prior to the burn for review and approval, and have the burn 

described in that prescription reviewed by the Division of Forestry during the burn operation. 

The local Division of Forestry District Manager (or their designee) will recommend DOF 

Prescribed Burn Manager certification upon satisfactory completion of both the prescription 

and required number of burns.

5. In order to continue to hold the Division of Forestry Prescribed Burn Manager Certification the 

burner must comply with 5I-2.006(2)(d), F.A.C., or Division Certification will terminate five 

years from the date of issue.

(d) Certification Renewal. A Certified Prescribed Burn Manager must satisfy the following requirements 

in order to retain certification.

1. Participation in a minimum of eight hours of Division of Forestry approved training every 

five years relating to the subject of prescribed fire, or participation in a Division of Forestry 

recognized Fire Council Meeting, and

2. The Certified Prescribed Burn Manager has submitted their certification number for two 

completed prescribed burns in the preceding five (5) years, or
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 255

3. Participation in five (5) burns and have this documented and verified in writing to the Forest 

Protection Bureau’s Prescribed Fire Manager of the Division of Forestry by a current Certified 

Prescribed Burn Manager, or

4. Retaking either the Prescribed Fire Correspondence Course or the Inter-Agency Basic Prescribed 

Fire Course.

(e) Decertification. The Commissioner of Agriculture will revoke any Certified Prescribed Burn 

Manager’s certification if they demonstrate that their practices and procedures repeatedly violated 

Florida law or agency rules or is a threat to public health, safety, or property. Recommendations for 

decertification by the Division of Forestry to the Commissioner of Agriculture will be based on the 

Certified Burner Violations—Point Assessment Table, effective July 1, 2003, which is incorporated by 

reference located at: http://www.fl-dof.com/wildfire/wf_pdfs/CBMpoints.pdf.
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Appendix C. Prescribed fire liability rules retained

on U.S. forest land by state (from Sun [in press])

Strict Liability Uncertain liability Simple negligence Gross negligence

Delaware Arizona Alabama Florida

Hawaii Colorado Alaska Georgia

Minnesota Connecticut Arkansas Michigan

Pennsylvania Idaho California Nevada

Rhode Island Illinois Kentucky

Wisconsin Indiana Louisiana

Iowa Maryland

Kansas Mississippi

Maine New Hampshire

Massachusetts New Jersey

Missouri New York

Montana North Carolina

Nebraska Oklahoma

New Mexico Oregon

North Dakota South Carolina

Ohio Texas

South Dakota Virginia

Tennessee Washington

Utah

Vermont

West Virginia

Wyoming

N =6 N =22 N =18 N =4
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 257

D. Sarasota County, Florida resolution supporting prescribed fire 

(on file with Sarasota County Board of County Commissioners)

RESOLUTION OF THE BOARD OF CUNTY COMMISSINERS

OF SARASOTA COUNTY, FLORIDA

RESOLUTION NO 97-265

RE: RESOLUTION IN SUPPORT OF PRESCRIBED BURNING

WHEREAS, The Environment Chapter of Apoxsee: The Revised and Updated Comprehensive Plan Policy 

5.5.10. states that Sarasota County is to “maintain and promote rural and natural resource land 

management practices such as prescribed burning;” and 

WHEREAS, The Recreation and Open Space Chapter of Apoxsee: The Revised and Updated 

Comprensive Plan Policy 1.2.5. states that “natural area parks,” once acquired, “will be kept in their 

natural state, receiving maintenance according to normal practice associated with native habitats;” 

and

WHEREAS, The Florida Legislature enacted Chapter 590.o26, Florida Statutes, that recognized the 

benefits of conducting prescribed burns; and

WHEREAS, prescribed burning is a critical resource management tool that when properly administered 

minimizes impacts on air quality, protects public safety and enhances scenic vistas; and

WHEREAS, a wide variety of ecosystems present within Sarasota County, many of which provide 

habitat for plants and animals classified as endangered, threatened or as species of special concern, 

require periodic burning to flourish; and

WHEREAS, prescribed burning reduces accumulated fuels and consequently lessens the likelihood and 

severity of uncontrolled and damaging wildfires; and 

WHEREAS, rules governing open burning in Sarasota County require that proper authorization be 

obtained form the Florida Division of Forestry, Department of Agriculture and Consumer Services 

and the Sarasota County Pollution Control Division prior to conducting prescribed burns in Sarasota 

County;

NOW THEREFORE BE IT RESOLVED by the BOARD OF COUNTY COMMISSIONERS OF 

SARASOTA COUNTY, FLORIDA, in public meeting assembled that;

The Board of County Commissions officially supports the use of prescribed burning as a land 

management tool in Sarasota County.
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Appendix E. Sarasota County Florida Comprehensive Land

Management Plan discussion of prescribed fire, and a

resultant “Notice of Proximity Conservation Easement” 

called Exhibit A in the U.S. 41/Blackburn Point Road Villade 

Activity Center Sector Plan No. 89-02-SP.which provides

smoke corridors through adjacent properties when using

prescription fire on designated State and County lands.

Environment Chapter

The Sarasota County Comprehensive Plan

Page 2-1

The Importance of Fire as a Management Tool

Very serious degradation of native plant and animal communities occurs when fire is suppressed in fire 

dependent habitats. Because frequent fire is essential to most native plant communities in Sarasota County, 

prescribed burning is now used, and has been used for generations by both preserve managers and ranchers to 

maintain pine flatwoods, scrub prairies and marshes. These plant communities are flammable by nature. When 

fire is intentionally excluded, fuels in the form of live and dead plant materials buildup, setting the stage for 

unintentional, often uncontrollable fires. Naturally occurring wildfires potentially threaten homes built in these 

areas. This is especially true as subdivisions are occurring in the rural and semi-rural areas.

Although residential destruction by wildfire in Florida is not as serious a problem as it is in western states, it is a 

reality. Since 1980, hundreds of Florida homes have been destroyed by wildfire. As in the recent history of North 

Port wildfires, these fires frequently start under conditions which are difficult to control; for example, during 

severe drought years. Prescribed ecological burning maintains native plant communities and reduces hazardous 

fuel accumulations.

Although an essential practice, prescribed burning must be regulated to minimize the impacts on surrounding 

land uses and residents. Burning impacts air quality as it releases particulates and carbon monoxide into the air. 

These impacts can be minimized through proper coordination with the Sarasota County Air Quality Monitoring 

Program which regulates open burning through Ordinance 72-37, as amended.

There is a growing conflict between rural land practices and suburban land uses. Recent development pressures 

in the rural areas of Sarasota County seriously threaten the continuance of prescribed burning and, thus, the 

maintenance of natural areas.

People who live and maintain farming operations in the rural areas may understand the role of fire in Florida’s 

natural areas and may be tolerant of controlled burning. However people from urban and suburban areas may 

be less tolerant of nearby fire and smoke. Environmental education efforts should help to inform the public 

that prescribed burning is part of the character of Southwest Florida and Sarasota County, and essential to 

maintaining native plant communities.

To encourage the practice of prescribed burning and minimize its impacts on surrounding land uses, the County 

could recognize the concept of smoke management buffer zones; especially in the eastern portions of the County 

near Myakka River State Park, the T. Mabry, Carlton, Jr. Memorial Reserve, and surrounding ranchlands. A 

suggested smoke management zone would extend approximately two miles from the boundaries of areas that 

depend on prescribed burning for land and wildlife management practices. After smoke management zones are 

outlined, all development proposals (rezoning, DRI, and special exceptions) within these zones would include 

acceptance of occasional smoke, a pre-existing condition or character of the area, as a condition for approval. 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 259

Care taken to plan compatible development in smoke management zones and implementation of 

the above policy will help insure the future of the County’s natural areas which are threatened by 

encroaching development restricting the necessary management practice of prescribed burning. As part 

of the management of conservation and preservation areas in urban and semi-rural settings, alternatives 

to prescribed fire should be researched and encouraged to maintain fire dependent communities.

EXHIBIT A

NOTICE OF PROXIMITY TO OSCAR SCHERER STATE

RECREATION AREA/CONSERVATION EASEMENT

This Notice date this ______ day of ______________________________, 199-, and entered into the 

public record by ____________________________________and ___________________________, 

as owners of the property described as:

SEE ATTACHED EXHIBIT I

(Insert description of subject property owned within U.S. 41/Blackburn Point Road Sector Plan No. 

89-02-SP)

WHEREAS, it is the intent of this Notice to make known to the public-at-large that the 

property described in Exhibit “I” attached hereto is located in close proximity to the property known as 

the Oscar Scherer State Recreation Area/Conservation Easement

WHEREAS, it is further the intent of this Notice to advise potential tenants and purchasers 

of subdivided property located within the boundaries of the property described in Exhibit “I” attached 

hereto, that said property is in close proximity to the Oscar Scherer State Recreation Area/Conservation 

Easement.

NOW, THEREFORE, the general public and those parties specifically purchasing or leasing 

property within the area described in Exhibit “I” attached hereto are hereby notified that:

 1. The subject property described in Exhibit “I” attached hereto is located in close proximity to 

the Oscar Scherer State Recreation Area/Conservation Easement.

 2. This Notice is to further advise potential purchasers or tenants of property described in 

Exhibit “I” attached hereto that the proximity to the Oscar Scherer State Recreation Area/Conservation 

Easement may result in said purchasers or tenants being affected by: continuing current resource 

management practices to include but not be limited to ecological burning, pesticide usage, exotic 

plant and animal removal, usage of heavy equipment and machinery and other practices as may be 

deemed necessary for the proper management of the Oscar Scherer State Recreation Area/Conservation 

Easement.

3. The nature and extent of the effects of the operations of the Oscar Scherer State Recreation 

Area which shall include: All management practices as contained within the document entitled 

“Ecological Burn Plan Oscar Scherer State Recreation Area” adopted on April 3, 1990, and which may 

be amended from time to time.
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4. All property owners which take title to property within the boundaries as described in Exibit 

“I” attached hereto, or tenants who may occupy the premises within the boundaries described in Exhibit 

“I” attached hereto, shall be deemed to have constructive knowledge of this Notice due to its recordation 

in the Public Records of Sarasota County, Florida, and further shall be deemed to have consented to 

said resource practices, including ecological burning, pesticide usage, exotic plant and animal removal, 

usage of heavy equipment and machinery and other practices as may be deemed necessary for the proper 

management of the Oscar Scherer State Recreation Area/Conservation Easement by the recording of a 

Warranty Deed or other instrument of conveyance, conveying the property within the boundaries in 

Exhibit “I” attached hereto, or by executing an occupancy agreement and delivering same to the owner 

of property contained within the boundaries of the property described in Exhibit “I”, their successors or 

assigns.

IN WITNESS WHEREOF, the owners have hereunto set their hands and seals this ____ day of 

_________________________________________________, 199-. 

_________________________________ _________________________________

_________________________________ _________________________________

STATE OF FLORIDA

COUNTY OF SARASOTA

I HEREBY CERTIFY that on this day before me, an office duly qualified to take 

acknowledgments, personally appeared __________________________________ and ____________

____________________, to me known to be the persons described in and who executed the foregoing 

instrument and acknowledged before me that they executed same. 

WITNESS my hand and official seal in the County and State last aforesaid this ____ day of 

________________________________________, 199-.

______________________________

NOTARY PUBLIC

 My Commission Expires:

(NOTARY SEAL)

093-007 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 261

Appendix F. Items to consider when drafting a statute to

protect and facilitate the intentional use of fire.

1. Require an authorization or permit to conduct any free-spreading outdoor fire. Require and keep

information on every requested burn including the purpose and size of the intended burn

2. Provide help in planning and executing burns

3. Provide standby during a burn as available

4. Clearly state that the intentional use of fire is a landowner right and, because it benefits the

environment, society in general, and the economy of the state, it can’t be declared a public nuisance

5. Authorize the state agency responsible for fire management (rather than the state agency responsible

for air quality) to take the lead in formulating and enforcing smoke management guidelines

6. Not place any arbitrary time of day, day of the week, or holiday restrictions on when burning can

occur. Such decisions should be based on measurable environmental conditions.

7. Develop a burner certification program with incentives that give such burners a higher level of

liability protection and more leeway in the execution of burns under their direct supervision. Also

develop re-certification procedures 

8. Consider tax incentives for ecosystems maintained with the judicious use of fire

9. Include language that provides personnel from other organizations and states assisting in fire

management operations at least the same level of protection their home organization/state provides

in case of an accident

10. Provide wildland fire suppression equipment and more training for Volunteer Fire Departments

11. Supplement equipment that is no longer available from pulp and paper companies

12. Enhance fire prevention and FireWise outreach efforts

13. Make it a punishable offence if a landowner is asked to, but refuses to manage his fuels and a fire

starts on his property, regardless of the cause, leaves his property, and damages the property of

others

14. State unequivocally that suppression forces do not have to attempt to suppress fires on property

where the owner has refused to manage fuel loads 

15. Invest in continued training for fire staff. Facilitate their attendance at training and refresher courses 

on agency time, even if out-of-state and reimburse them for expenses 

16. Provide funds to keep fire equipment up-to-date and well-maintained.

17. Provide personal protective gear for all fire staff and include language to make all other state 

agencies with fire staff to do the same. Make its use mandatory on all fires

18. Provide funds to enter into an agreement to have a state meteorologist provide daily fire danger 

forecasts and timely spot weather forecasts for complex burns upon request. Require and fund 

training and expose them to operational fires so they can hone their skills. Several southern state 

forestry agencies have their own forest meteorologist, or pay a neighboring state that has one, to 

handle their fire meteorology duties
2	 Fire 62 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

19. Include language to try to protect fire staff and scheduled equipment replacement during tight 

budgets. Clearly identify fire staff as 1st responders in natural disasters and other emergencies. Train 

all fire staff on the Incident Command System (ICS) and require its use statewide in emergencies

20. Facilitate formation of a fire council, fund development and maintenance of a web site for it, 

encourage fire staff to attend and assume leadership roles on agency time, and reimburse them 

when they do. 

21. Encourage the development of incentives to retain experience and to reward people who 

successfully accept these risks associated with burning.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 263

Growing a Burning Program: Challenges and Opportunities

Rex Mann1

...there is much ground cleared by the Indians, and 

especially about (their agricultural fields); and I 

am told that about three miles from us a man may 

stand on a hilly place and see thousands of acres of 

ground as good as need be, and not a Tree on the 

same.

In 1637, Thomas Morton wrote that the Indians:

...are accustomed to set fire of the Country in all 

places where they come, and to burne it twize, in 

the year...as the Spring and fall of the leafe...so 

that hee that will looke to find large trees and good 

tymber...(will not) finde them on upland ground; 

but must seeke for them...in the lower grounds, 

where the grounds are wett.

But frequent forest burning did more than reduce the 

undergrowth and improve the habitat for preferred 

species. In many cases it created grasslands in areas where 

forest otherwise would have existed. Prairies extended 

into Ohio, western Pennsylvania, and western New 

York. In Virginia, the Shenandoah Valley—a broad 

valley located between the Blue Ridge Mountains and 

the Allegheny’s—was on vast grass prairie that covered 

more than 1,000 square miles. Native Americans burned 

the area annually. R.C. Anderson wrote that the eastern 

prairies and grasslands “would mostly have disappeared 

if it had not been for the nearly annual burning of these 

grasslands by the North American Indians.” In the West, 

as well, Indian burning greatly extended the area of 

grasslands and reduced the area of forest.

Exactly why the Indians burned is less clear. Most likely 

they used fire, as did early peoples on all continents, to 

shape and affect landscapes in ways that made it easier 

for them to live. Fire was the tool that cleared land for 

agriculture and created conditions favorable for deer 

and other wildlife. This included creating patches of 

blueberries as well as conditions that favored oaks and 

chestnuts. In other words, the use of fire was closely 

related to obtaining food. Since the Indians had little use 

for hardwood sawlogs, you have to believe that firing the 

Let me emphasize that I only claim to speak with 

authority about the task of restoring fire to the National 

Forests. Private landowners have their own special 

problems in using prescribed fire, size of forest tracts, 

lack of expertise, etc.

Before we can appreciate the challenges we face in 

returning fire to the oak forests of the Eastern United 

States, we must understand the old paradigm that says 

that “all fire in the forest is bad.” We need to understand 

how the old paradigm came to be, how it was reinforced 

over the years, and how it still persists in the minds of 

many if not most of our citizens.

The fact that we had this conference illustrates the 

change in thinking of land managers and the scientific 

community on the proper role of fire in our oakdominated forests. If we could travel back in time to the 

end of the 19th century and the early years of the 20th 

century, we might hear debates between fire advocates 

and fire opponents that sound much like those going on 

today.

There is ample evidence from early journals, letters, and 

other historical documents that the European settlers saw 

abundant use of fire by Native Americans. For example, 

John Smith commented that in the forests around 

Jamestown, Virginia, “a man may gallop a horse amongst 

these woods any waiy, but where the creeks and rivers 

shall hinder.” Andrew White, on an expedition along 

the Potomac in 1633, observed that the forest was “not 

choked with an undergrowth of brambles and bushes, 

but as if laid out in a manner so open, that you might 

freely drive a four house chariot in the midst of the 

trees.”

In 1630, Francis Higginson wrote about the country 

around Salem, Massachusetts, that:

1

Emergency Management Specialist, USDA Forest Service, 

1700 Bypass Road, Winchester, KY 40391, 859-745-3123, 

email: rmann@fs.fed.us.
2	 Fire 64 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

woods was considered an act of stewardship rather than 

an act of destruction.

There also is ample evidence that early Europeans 

continued the custom of burning the forest, long after 

the Indian populations had collapsed from exposure 

to our diseases. This burning was mostly to promote 

grazing for livestock, including hogs.

In the closing years of the 19th century, the Nation 

was experiencing an unprecedented wave of destructive 

logging and land clearing. As Stephen Pyne expressed in 

“Year of the Fire”: “Above all, fire–abusive fire—followed 

the ax, flames fed on the extravagant wreckage left by 

logging and land clearing.” Steam engines, used in 

logging, provided sparks for easy ignition of logging 

slash. Destructive fires occurred in places that seldom 

have fire problems today, such as New England and New 

York. Entire towns and villages burned, with significant 

loss of life. Concurrent concerns about the liquidation of 

the nation’s forests and the widespread fires set the stage 

for the creation of the USDA Forest Service in 1905 

and the establishment of state forestry agencies in the 

immediate decades that followed.

Our early foresters were educated in Europe, the one 

place where agriculture and settlement had changed 

landscapes to the point where fire no longer played a 

natural role. They brought to America the view that fire 

had no place in the forest. As early as the 1809’s, Bernard 

Fernow, the Prussian trained forester who headed the 

Division of Forestry, had denounced America’s ceaseless 

burning as a product of “bad habits and loose morals.” 

Gifford Pinchot, who more than anyone else shaped 

American forestry, had this to say: “The question of 

forest fires, like the question of slavery, may be shelved 

for a time, at enormous cost in the end, but sooner or 

later it must be faced.”

There were voices arguing for continuing the “light 

burning” or Indian burning that had been practiced 

for centuries. Major John Wesley Powell, director of 

the U.S. Geological Survey, had come to believe that 

the practice of regular burning was a surer method of 

protecting the land from wildfire than attempting to 

abolish fire altogether, William B. Greeley Associate 

Chief of the Forest Service, contemptuously referred to 

this philosophy as “Paiute forestry.”

Then came the fires of 1910, called the Big Blowup, 

when millions of acres burned and scores of firefighters 

died. Although largely started from steam engines and 

logging slash, these fires solidified the Forest Service’s 

position that fire had no place in American forestry. The 

state forestry agencies and the Department of the Interior 

also adopted the “all fire is bad” paradigm.

This position on fire was maintained for decades, 

strengthened by Smokey Bear during World War II. It 

was only in the piney wood of the South that burning 

became an accepted practice.

By the end of the 20th century, many of the public lands 

suffered fire famine. Forests were diseased and dying and 

prone to catastrophic fire. In the East, species such as the 

red cockaded woodpecker were becoming more rare and 

species such as the American chaffseed were disappearing. 

The uplands had been invaded by fire intolerant species 

that likely had never grown there before.

American society, no longer rural, had shed its familiarity 

with fire and now feared it. Urban values had replaced 

rural values: fire was regarded as an enemy and not a 

useful tool or a vital process.

So here we are—a century after the Forest Service was 

formed, a century after the American public was told 

that fire in the forest was bad, a century after those 

who argued for “light burning” were ridiculed and 

dismissed—faced with the need to explain to the public 

that the founding foresters were wrong, that we now 

know that fire plays a vital role in sustaining our oak 

forests.

Of course, hindsight always is perfect. This epiphany 

that fire is now useful did not occur overnight. A good 

deal of research has been and is being conducted on 

the proper role of fire in our forests. Some of the most 

significant research has been paleo-botanical, where the 

stories told by ancient grains and charcoal fragments 

challenged the basic tenets of land managers who for 

decades had excluded fire.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 265

On the national forests we have finally learned that 

whatever management program we undertake must have 

public support. We can do the burning but how do we 

gain that support?

What are our challenges?

1. Our public is now largely urban and in urban 

areas fire generally is considered an enemy or a 

bad thing.

2. The public had little knowledge of fire as a tool, 

farmers burning fields or pastures, burning 

brush piles, etc.

3. Our TV sets give us round-the-clock images of 

horrendous wildfires each summer that burn 

homes and kill people.

4. We have had some spectacular prescribed fire 

escapes, beginning with Mack Lake in 1980 

on the Huron-Manistee National Forest in 

Michigan. This fire burned 20,000 acres and 

44 homes and buildings; one firefighter was 

killed. The Las Alamos fires in 2000 destroyed 

235 homes and triggered one of the largest and 

most expensive post-fire rehabilitation efforts 

in history. Recently, an escaped fire in Florida 

burned 38,000 acres.

5. The smoke from prescribed fires is ours no 

matter where it goes. Unlike the Indian burning 

of centuries ago, we must be concerned about 

where our smoke goes.

6. There are some who believe lightening fire is 

good, but human-ignited fire is bad. This belief 

is rooted in the myths of the pristine forest and 

the ecologically invisible Indian, in the mistaken 

belief that North America’s First People could 

not or would not use fire to shape landscapes.

7. There are some who believe that scarring the 

base of a sawlog tree is morally wrong. Given 

the variable fire effects we can expect when we 

install a fire regime in the oak uplands, we know 

we can strive to minimize this type of damage, 

yet it is unreasonable to believe we can totally 

eliminate it.

Here are additional challenges:

1. Will our workforce and budgets allow us to 

grow this program to the required level?

2. Do we have internal resistance to burning 

on this scale? Is there significant opposition 

from Forest Service employees and/or our 

cooperating agencies?

3. How do we get around the limited burning 

days we have? Of all of the factors that limit 

our ability to grow a program, the dozen or so 

suitable burning days annually seem to be the 

most severe.

Last year on the Daniel Boone National Forest, we 

successfully prescribe burned more acreage than ever 

before—a little more than 19,000 acres. Our new forest 

plan calls for us to burn 50,000 acres annually by the 

end of the next decade. We intend to grow this program 

carefully but steadily by several thousand acres each 

year. On a forest of more than 700,000 acres, 50,000 

acres burned per year on a 3- or 4-year cycle probably 

is inadequate for restoring and sustaining the forest 

conditions we are seeking on our uplands.

As stated earlier, of equal importance to growing the 

program is gaining public support or acceptance for 

the program. We intend to do this by reaching out 

to our publics to participate in the actual planning 

of landscape projects to promote a healthier forest 

condition. This is a critical point. Our new forest 

plan is driven by a concern for forest health, not how 

many board feet of lumber we can produce or how 

many acres we can burn. On the uplands of the Daniel 

Boone we will be creating a variety of forest conditions, 

such as pine and hardwood grassland communities, 

woodlands, and other habitat types that reduce the 

overcrowded conditions resulting from 70+ years of fire 

exclusion. These communities will be treated carefully. 

Commercial logging will be used to reduce the stocking 

and remove fire-sensitive species that have invaded the 

sites. To the degree possible, lands not immediately put 

into these special conditions will be thinned to reduce 

overcrowding.
2	 Fire 66 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Fire will be reintroduced into these uplands not as an 

end in itself but as a vital process that maintains the 

healthier conditions we have placed on the landscape. 

We are not viewing the reintroduction of fire as a single 

event, or as several successive burns, but as a permanent 

recurring process that is vital to sustaining what we 

believe are healthier, more resilient systems. Along with 

restoring fire, we intend to restore the American chestnut 

in partnership with the American Chestnut Foundation, 

and, hopefully, other lost species. I should say also that 

our goal is not to eliminate red maple, white pine, or 

hemlock but to confine these species to the wetter stream 

bottoms where they historically grew.

We are fully aware that this will not be an easy sell. We 

also are aware that much of the forest will not be burned, 

because of what I call social constraints e.g., interstate 

highways, airports, hospitals, nursing homes, and 

recreation areas, and our fear of affecting these areas with 

smoke.

We must educate the public as we involve it. It is critical 

that we speak of fire as a vital process in a larger effort of 

creating and sustaining healthier forest conditions. We 

must discard the old “all fire is bad” paradigm and help 

the public understand that, like wind and rain, fire is a 

vital, natural process.

At the implementation level, there are a number of 

things we must do to grow this program. We are 

limited by the number of burning days that occur. This 

means that on a suitable burning day, smoke must rise. 

Prescribed fire is not necessarily more important than the 

other jobs we do, but it is time sensitive. We must burn 

large areas; at least 500 acres in size It takes a day to burn 

a hundred acres or a thousand acres. We must use aerial 

ignition because it is so much faster and cheaper. Using 

ping-pong ball dispensers, we can ignite 1,000 acres 

in less than an hour. We must consider fall and winter 

burning and summer burning when our new habitat 

conditions are in place with a changed fuelbed. We must 

learn to ignite fire and fight fire at the same time—in 

Kentucky for example, arson fires are perhaps the greatest 

obstacles to growing a program. We must investigate the 

possibility of nighttime burning to extend our burning 

window.

We must do all this without succumbing to the urge 

to burn on days when conditions are too severe. The 

recent history of escaped burns teaches us that the public 

acceptance of controlled burning is fragile and disastrous 

escapes can kill a program.

Regardless of the challenges and obstacles, we must 

move wisely but aggressively to reintroduce fire to the 

uplands of the Daniel Boone. We recently lost most of 

our southern yellow pine trees and the northern-most 

group of red cockaded woodpeckers to a native insect, 

the Southern Pine Beetle. A major factor in this loss was 

the overcrowded, stressed conditions of this ecosystem, 

which had developed after decades of fire exclusion. The 

same disaster could lie in store for us in our hardwood 

uplands if we fail to act.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 267

Poster Abstracts
2	 Fire 68 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

OVERSTORY AND SEEDLING RESPONSE TO REPEATED FIRES IN 

EASTERN KENTUCKY

Heather D. Alexander1†, Mary Arthur2, David Loftis3, and Stephanie Green4

Oak (Quercus spp.) regeneration failure throughout eastern deciduous forests has prompted forest 

managers to use prescribed fire as a potential tool to maintain oak communities. Within the Daniel 

Boone National Forest of eastern Kentucky, we tested the ability of repeated prescribed fires to 

increase oak seedling growth and survival by increasing light at the forest floor, decreasing competitor 

seedling survival, and enhancing oak seedling leaf characteristics. We measured stand structure, 

canopy cover, seedling survival, and seedling leaf characteristics in areas unburned, burned once, and 

burned twice. Pre-burn density measurements showed oaks dominated overstory classes but maples 

(Acer spp.) and other species dominated the midstory and understory. In areas burned twice, canopy 

cover temporarily decreased after the first fire but quickly returned to pre-burn levels after the second 

fire due to increased sprout density. Burning did not affect red oak (Erythrobalanus) survival, but 

white oak (Leucobalanus) survival decreased. Among competitor seedlings, burning had no effect on 

sassafras (Sassafras albidum (Nutt.) Nees)) survival, but maples (red and sugar maple combined) had 

the greatest mortality of all species. Within burned areas, competitor seedlings produced more leaves 

with greater leaf area than red or white oaks. Burning led to higher specific leaf mass (leaf thickness 

per unit area) for both oak groups compared to competitor seedlings; this increase was correlated with 

increased light levels. Repeated fires temporarily opened up the canopy but post-fire sprouting rapidly 

cancelled this positive effect. Burning also reduced the survival of red maple seedlings, but competitor 

seedlings that survived the fire grew more leaves and had higher leaf area than oaks. Burned oaks had 

thicker leaves but whether this leads to increased growth is unknown. These findings suggest multiple, 

repeated fires with short burn intervals may be needed to maintain canopy cover and sufficient 

understory competition to successfully regenerate oaks.

1

Ph.D. Candidate, University of Kentucky, Forestry Department; 2

Associate Professor, University of Kentucky, Forestry Department; 3

Project Leader, USDA Forest Service, Bent Creek Experimental Forest; 4

Senior Land Management Specialist, Southwest Florida Water 

Management District; †Corresponding author, T.P. Cooper Bldg., 

Lexington, KY 40546, 859-257-2806, email: heather.alexander@uky.edu
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 269

THERMOCOUPLE PROBES IN SURFACE FIRES

Anthony S. Bova

1†

 and Matthew B. Dickinson2

The maximum temperatures of thermocouple probes (TCPs) exposed to wildland flames often are 

called “fire temperatures” in forest fire literature, but such temperatures depend on TCP diameter, 

configuration, and physical properties, as well as flame temperature, dimensions, and velocity over the 

TCP surface. Thus, TCP temperatures alone give no useful information about a surface fire. However, 

in small experiments, TCP response was calibrated to provide general fire characteristics. For 

instance, we found that the maximum time-rate of change of TCP temperature and the area under 

the temperature-time curve correspond to the fireline intensity and fuel consumption, respectively. 

Calibration equations derived from these data can be used to estimate fireline intensity and fuel 

consumption from multiple TCPs installed at remote locations in prescribed burns.

1

Physical Scientist and 2Ecologist, Northern Research Station; †

Corresponding author, 359 Main Road, Delaware, OH 43015, 

740-368-0093, email: abova@fs.fed.us.
2	 Fire 70 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Wildfire Hazard Mitigation Treatments Applied Above ground

Impact Soil ecological properties in Oak Forest Ecosystems

Jennifer A. Brinkman1† and R.E.J. Boerner2

Manipulations designed to reduce wildfire hazard in forests in which wildfire has been suppressed 

for many decades typically entails manipulating fuels with prescribed fire and mechanical treatment. 

Although the ecosystem properties being manipulated are above ground, these activities also may 

affect below-ground components of ecosystems. We evaluated soil organic matter and microbial 

activity in mixed-oak (Quercus spp.) forests of southern Ohio that were subjected to alternative 

wildfire hazard modalities (prescribed fire, thinning from below, and the combination of thinning and 

burning) and untreated controls over a 4-year period. During the first growing season after treatment, 

soil organic C quantity was significantly lower in soil from the thin+burn plots than from the other 

three treatments, but there was no significant difference in organic matter quality (C:N ratio) among 

treatments. By the fourth year after treatment, the soils from the thinned plots had significantly 

greater organic matter quantity and lower organic matter quality than soils from the other treatments. 

During the first year, net N mineralization increased after thinning and decreased after fire relative 

to the untreated control. Plots that were both thinned and burned did not differ from the control. 

By the fourth year, there were no differences among treatments in N mineralization. Proportional 

nitrification was significantly lower in thinned plots (with or without fire) than in control or burned 

plots in the first year, but this pattern was reversed by the fourth year. Acid phosphatase activity (an 

index of microbial breakdown of labile carbon compounds) was lower in the thin+burn plots that 

in the other three treatments the first year, but by the fourth year, acid phosphatase activity was 

greater in the thinned plots than in the other treatments. Phenol oxidase activity (an index of fungal 

breakdown of recalcitrant organic matter) was not significantly affected by any treatment in the first 

or fourth year after treatment. The responses to the hazard reduction treatments differed among 

response parameters and years; thus, ordination approaches were used to present a more holistic view 

of the impact of above-ground manipulations forest-soil ecology.

1

Research Assistant, The Ohio State University, Department of 

Evolution, Ecology, and Organismal Biology; 2Professor, The Ohio State 

University, Department of Evolution, Ecology, and Organismal Biology; †

Corresponding author, 318 W. 12th, 300 Aronoff Laboratory, Columbus, 

OH 43210 USA, 614-292-1688, email: brinkman.3@osu.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 271

FIRE RESEARCH IN THE PINE BARRENS OF NEW JERSEY

Kenneth Clark1†, John Hom1, Nick Skowronski1, Yude Pan1, Steve Van Tuyl1, and Warren Heilman2

We are developing an interdisciplinary fire research program based at the Silas Little Experimental 

Forest in the Pine Barrens of New Jersey to better estimate wildfire risk, quantify the tradeoffs 

between hazardous fuel reduction treatments and carbon sequestration by forests, and measure 

the impact of hazardous fuel reduction on emissions of EPA-criterion pollutants near urban 

nonattainment areas. We are using an integrated network of field plots, fire weather and eddy flux 

towers, remotely sensed data layers, and validated fire weather and forest ecosystem models. Extensive 

field plots and LIDAR measurements have been used to estimate fuel loading and the presence of 

ladder fuels across the Pinelands. In addition, pre- and post-prescribed fire measurements of fuel 

loading are used to evaluate fuel reduction treatments. These activities provide better estimates of 

hazardous fuel loading and the cost effectiveness of fuel reduction treatments in the Pine Barrens. We 

installed and operate six new fire weather towers providing real-time weather data to fire managers 

over the internet (http://climate.rutgers.edu/stateclim/). A fuel moisture index based on forest energy 

balance measurements made from the flux towers is operational in New Jersey, and is being tested at 

a number of Ameriflux sites. We are using the network of fire weather and flux towers in conjunction 

with a SODAR to measure windspeed and direction up to a height of 700 m to validate predictions 

of MM5, a regional fire weather model, and smoke emission models. Biometric measurements, 

flux towers, and ecosystem- to landscape-scale models (PnET CN, BiomBGC) are used to estimate 

rates of forest productivity and fuel accumulation in the Pine Barrens. Collectively, these data are 

leading to the “next generation” of fuel models in which the dynamics of 1, 10, and 100 fuels and 

fuel-reduction treatments are integrated to estimate transitions among fuel models in a GIS database. 

These research products and other decision-support tools for managers and policymakers enable the 

latest science-based knowledge to be incorporated into decision processes at local to regional scales.

1† USDA Forest Service, Northern Research Station, Northern Global 

Change Program, Newtown Square, PA 19073, 2Northern Research 

Station, USDA Forest Service, East Lansing, MI 48823 †Corresponding 

author; 609-894-0325, email: kennethclark@fs.fed.us.
2	 Fire 72 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

EFFECTS OF FUEL REDUCTION TREATMENTS ON SOIL NITROGEN 

TRANSFORMATION IN THE SOUTHEASTERN PIEDMONT

T. Adam Coates1†, Victor B. Shelburne2, Thomas A. Waldrop3, Bill R. Smith4, and Hoke S. Hill, Jr.5

The Clemson Experimental Forest (CEF) in northwestern South Carolina is one of 13 sites included 

in the National Fire and Fire Surrogate Study. Fuel-reduction treatments (prescribed fire, thinning, 

and a combination of the two) were installed on the CEF beginning in the winter of 2000-2001. The 

inorganic nitrogen (N) contents of the soil (KCl extractable NH4

+

-N and NO

3

-

-N) were determined 

prior to treatment and in 2004 and were used to calculate three measures of soil N transformation: 

the rate of mineralization, the rate of nitrification, and proportional nitrification. Using an analysis 

of covariance, it was determined that the 2004 results for N transformation were not significantly 

influenced by the levels of N transformation before treatment. Analysis of variance for the 2004 

results showed no significant differences among treatment means, and linear contrasts for these 

data suggest that none of the fuel reduction treatment means differed significantly from the control 

means. Differences before and after treatment were not significant for all variables when paired ttests were used. The nitrification rate increased following these treatments in the thin only and burn 

only plots. These analyses suggest that the treatments have had no significant detrimental effect on 

soil N transformation over the 2- to 3.5-year posttreatment period, and may increase the rate of 

nitrification.

1

Graduate Student, Department of Forestry and Natural Resources, 

Clemson University; 2Professor, Department of Forestry and Natural 

Resources, Clemson University; 3Research Forester, USDA Forest Service, 

Southern Research Station; 4Professor, Department of Entomology, Soils, 

and Plant Sciences, Clemson University; 5Department Chair, Department 

of Applied Economics and Statistics, Clemson University; †Corresponding 

author, 265 Lehotsky Hall, Clemson, SC 29634, 864-656-3054, email: 

coatest@clemson.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 273

Use of Wildland Fire IN Great Smoky Mountains National Park

Dana Cohen1†, Beth Buchanan2, Bob Dellinger3, Leon Konz4, Mark Taylor5

Great Smoky Mountains National Park, located in Tennessee and North Carolina, had a policy 

of suppressing all fires from its inception in 1934 until the adoption of wildland fire use policy in 

its 1996 Fire Management Plan. This enables park fire staff to manage lightning-caused fires to 

accomplish resource objectives so long as the fires meet certain predefined conditions. The Smokies 

was the first federal land management unit in the Southern Appalachians to use naturally ignited 

fires. Unlike prescribed fires, lightning-caused fires tend to burn over long periods with a variety 

of behavior and effects in a mosaic over the landscape. They have been an essential tool for better 

understanding the natural role of fire in the park.

1

Fuels Specialist, Dixie National Forest; 2Fire Ecologist, USDA Forest 

Service, Southern Region; 3Biological Science Technician (Fire Effects), 

North Cascades National Park; 4Fire Management Officer (ret.), Great 

Smoky Mountains National Park; 5Fire Management Officer, Great 

Smoky Mountains National Park; †Corresponding author, P.O. Box 645, 

Panguitch, UT 84759, (435) 676-9372, email: dcohen@fs.fed.us.
2	 Fire 74 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

HARDWOOD STEM INJURY AND MORTALITY IN SURFACE FIRES

Matthew B. Dickinson1† and Anthony S. Bova

2

In surface fires, hardwood stems are heated primarily by flames.When the heat flux to those stems is 

sufficiently great or prolonged, necrosis of phloem and live sapwood tissues results. We are developing 

models of hardwood stem injury and mortality along several fronts. In collaboration with the 

Missoula Fire Sciences Laboratory, we are developing a stem injury and mortality model. A version 

of this model designed for fire managers is called FireStem (www.firelab.org). Estimating heat flux 

to stems is critical as an input to any stem heating model, but accurate estimates are difficult and 

expensive to obtain. To address this problem, we are developing an inverse heating methodology to 

estimate net stem heat flux that requires minimal instrumentation and has the additional advantage 

of using the tree stem itself as a sort of heat flux sensor. Once stem heating can be predicted, it is 

necessary to have a tissue necrosis model. Two approaches are being pursued for developing predictive 

models of tissue necrosis. In one, equations whose form is determined by heat transfer principles are 

parameterized by data on fire behavior, heat flux, and tissue necrosis. In a more biophysically realistic 

approach (that used in FireStem), tissue necrosis models are linked with the stem heating model. 

For small stems, heating may be roughly equal around stems, but uneven heating is the general 

result where fires burn on slopes or in wind. Field data and modeling are being used to improve our 

understanding of uneven heating so that it can be incorporated in stem injury and mortality models.

1

Ecologist and 2Physical Scientist, USDA Forest Service, Northern Research 

Station; †Corresponding author, 359 Main Road, Delaware, OH 43015, 

740-368-0093, email: mbdickinson@fs.fed.us.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 275

vegetation response to prescribed fire across a moisture/

productivity gradient in the southern appalachians

Katherine J. Elliott† and James M. Vose

We examined the effects of single, dormant-season fires on overstory, midstory, and ground flora 

species diversity and composition, overstory mortality, and tree seedling regeneration the first and 

second years following prescribed burns in eastern Tennessee, northern Georgia, and western North 

Carolina. We evaluated the effects of burning along a moisture/productivity gradient that ranged 

from dry, pine-hardwood to mesic, mixed-hardwood ecosystems. All treated sites were burned in 

spring before leaf-out (March-April), and adjacent sites were designated as controls. Within each site, 

vegetation plots were measured in layers: the overstory layer (trees ≥ 5.0 cm d.b.h.); the midstory 

layer (woody stems < 5.0 cm d.b.h. and ≥ 0.5 m height); and the ground flora layer (woody stems 

< 0.5 m height and all herbaceous species). Permanent plots were sampled before the prescribed burns 

and the first and second growing seasons after the burns. As a result of the fires, overstory mortality 

occurred primarily in the small size-class hardwoods and ranged from 23 percent in the dry, pinehardwoods forest to < 2 percent in the mesic, mixed-hardwoods forest. In the midstory, fire reduced 

the basal area of woody species but prolific sprouting from hardwoods resulted in higher density 

of fire-sensitive hardwoods such as red maple (Acer rubrum L.), sourwood (Oxydendrum arboretum 

(L.) DC.)), and blackgum (Nyssa sylvatica Marshall). On the dry sites, blueberries (Vaccinium spp.), 

huckleberry (Gaylussacia ursina (M.A. Curtis) T & G)), and mountain laurel (Kalmia latifolia L.) 

increased in density. White pine (Pinus strobus L.) decreased in density by 20 to 50 percent and its 

basal area was reduced by 50 percent after burning. On the most mesic site, midstory density and 

basal area were not significantly changed after burning. We found no significant change in groundflora percent cover or diversity after prescribed on mesic sites. We hypothesized that the ground flora 

was not significantly affected by these low- to moderate-intensity fires because the burns occurred 

in the spring before the emergence of herbaceous species, and the low soil depth of heat penetration 

(< 2.0 cm at 59o

C) did not result in damage to below-ground rhizomes or buried seeds.

USDA Forest Service, Southern Research Station, Coweeta Hydrologic 

Laboratory; †Corresponding author, 3160 Coweeta Lab Rd, Otto, 

NC 28763, 828-524-2128, email: kelliott@fs.fed.us.
2	 Fire 76 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

OAKS IN PRESETTLEMENT FIRE LANDSCAPES OF THE SOUTHEAST

Cecil Frost1

Oaks are found in all physiographic provinces of the Southeastern United States and occurred in 

presettlement times at all fire frequencies, including 2- to 3-year fire intervals in the most fire-exposed 

landscapes. Both vegetation structure and species diversity have been altered greatly by some 70 

years of fire suppression. In the Sandhills of North and South Carolina, Michaux (1805) reported 

that “Seven-tenths of the country are covered with pines of one species, or Pinus palustris.” However, 

there was a variety of oak vegetation types within the pine-dominated southeastern landscape. The 

original percentage of upland oaks in one flat, south Atlantic Coastal Plain landscape was estimated 

at less than 1, 2.3 in a gently rolling site in the North Carolina Sandhills, and 15 at a more dissected 

site in South Carolina near the Sandhills-Piedmont transition. On the Piedmont itself was a band 

of post oak savanna in South Carolina and Georgia that ran parallel to and just west of the margin 

of the range of longleaf pine. With only a slightly lower fire frequency, about 2- to 5 years, this 

type consisted of an open canopy of post oak over a species-rich grassy herb layer. Habitats on the 

Southeastern Piedmont were more nearly balanced between pines—principally shortleaf pine—and 

hardwoods, especially white oak, post oak, mockernut hickory, red oak, and chestnut oak. In the 

Southern Appalachians, oak habitats were more varied and fire frequencies ranged from nearly annual 

in areas where the fire regime was dominated by Native American ignitions to about 5 years in some 

chestnut oak forests, to essentially fire free areas in cove hardwood stands where species such as white 

oak persisted through gap-phase succession rather than through regeneration by fire. In those stands 

where reproduction depended on fire, loss or depauperization of the herb layer in fire-suppressed oak 

forests is one of the most significant effects of fire suppression.

1

Landscape Fire Ecologist, North Carolina Plant Conservation Program 

(ret.), 119 Potluck Farm Road, Rougemont, NC 27572, 336-364-1924, 

email: cecil.frost@earthlink.net.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 277

EFFECTS OF PRESCRIBED BURNING ON INVASIBILITY BY NONNATIVE

PLANT SPECIES IN THE CENTRAL HARDWOODS REGION

Lance Glasgow and Glenn Matlack1

Fire often promotes invasion by nonnative plant species, yet few studies have examined this process 

in forests of the Central Hardwood Region. Public agencies are considering prescribed fire as a 

management tool to favor the regeneration of oak (Quercus spp.) in this region, raising the possibility 

of widespread invasion of forests by nonnative species. To examine the potential role of burning in 

facilitating invasions, we set up an experiment simulating various aspects of forest fires at a forested 

site in southeastern Ohio. Treatments included high- and low-burn intensity, increased soil pH, litter 

removal, and a control. Seeds of two problematic nonnative species (Microstegium vimineum and 

Rosa multiflora) were sown experimentally following application of the treatments. Treatments were 

arranged in a randomized block design in two landscape positions (dry upland and moist lowland) 

and two canopy conditions (gap, no gap). We measured germination, stem height, leaf number, 

and seed production at four time intervals throughout the growing season. Germination rate was 

promoted by litter removal, and high- and low-intensity fire treatments for M. vimineum, and by 

high-intensity fire for R. multiflora. In both species, stem height and leaf number were greatest 

following high-intensity fire and under canopy gaps. We suggest that prescribed burning creates 

a disturbance suitable to invasion. Removal of leaf litter seems to be the most important factor 

controlling germination of our experimental species, while overall site quality (light and moisture) 

is most important for seedling survival. Growth and reproduction of both species were highest in 

the hot burn treatments, indicating that the most severe disturbances have the greatest effect on the 

invasibility of deciduous forests. We infer that the absence of invasion in previous studies is due to a 

lack of propagules rather than the unsuitability of the burned site for germination or growth.

1

Environmental and Plant Biology, Porter Hall, Ohio University, Athens, 

Ohio, 45701, USA 740-593-1131; email Matlack@ohio.edu.
2	 Fire 78 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

FUEL AND FIRE DYNAMICS IN EASTERN MIXED-OAK FORESTS

John B. Graham1 and Brian C. McCarthy2†

In eastern mixed-oak forests, prescribed fire and thinning are being used as tools for a variety of 

management goals, including altering fuel loads. However, the specific effects of these treatments 

on fuel dynamics in hardwood forests are relatively unknown, as are the specific patterns of fire 

behavior based on fuel loading and moisture content. We conducted two experiments to address these 

deficiencies. First, we conducted a study to examine fuel dynamics 3 years after thinning, prescribed 

fire, and a combined treatment as a part of the Ohio Hills site of the Fire and Fire Surrogate program. 

Measurements techniques followed the Brown planar-intersect method to determine litter, duff, 1-hr 

(0 to 6 mm diameter), 10-hr (6 to 25 mm), 100-hr (25 to 75 mm), and sound (1,000S) or rotten 

(1,000R) 1,000-hr (75+ mm) fuel mass. Coarse woody debris (CWD) was evaluated for species, 

length, large- and small-end diameters, and decay class. Three years after treatments, thinning 

increased 100-hr, 1,000S, CWD, and litter, but reduced 1-hr fuels; burning increased 1,000S but 

reduced 1,000R; thinning followed by burning increased 100-hr, 1,000S, and CWD but reduced 

1-hr, 10-hr, and duff. Changes in the larger, sound fuels (100-hr, 1000S, and CWD) appear to 

persist over time following treatments, while changes in finer fuels (litter, duff, 1-hr, and 10-hr) or 

less-sound fuels (1,000R) appear to be more ephemeral, recovering or shifting direction within 3 

years after treatment. In the second experiment, we manipulated moisture and fuel-load levels (litter, 

1-hr, and 10-hr) in a common garden, trial burn experiment using fuel load estimates covering the 

range estimated in the first portion of the study. As predicted, fires burned hotter and consumed 

more fuel with elevated fuel loads or lower fuel moistures. The results of this experiment can be used 

by managers to predict fire behavior based on fuel conditions, and can be linked to other research 

to predict stem mortality based on fire temperature. Collectively, our experiments can be used to 

evaluate the effects of silvicultural treatments on future prescribed fires. As such, they provide a 

realistic baseline for fuel loading and fire behavior in models of eastern mixed-oak forests.

1

Graduate student, Ohio University, Dept. of Environmental and 

Plant Biology; 2Professor of Forest Ecology, Ohio University, Dept. of 

Environmental and Plant Biology; †Corresponding author, 315 Porter 

Hall, Ohio University, Athens, OH 45701, 740-593-1615, email: 

mccarthy@ohio.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 279

USING PRESCRIBED FIRE AND MECHANICAL THINNING TO IMPROVE

THE REGENERATIVE POTENTIAL OF OAK SPECIES

Scott Holevoet1† and Charles Ruffner2

Historical evidence and ecophysiological characteristics indicate a strong link between disturbance 

regime and the occurrence of oak (Quercus) species. Research indicates that many current oak 

forests developed following disturbances such as logging, fire, or a combination of logging and fire 

from the 18th century to the early 20th century. However, during the early and mid-20th century, 

many oak forests were entered for the last time and fire and cutting ceased. This resulted in the 

development of several cohorts of mixed mesophytic species, which are poised to replace the overstory 

with the next disturbance. To counter this, managers are being encouraged to use prescribed fire 

to improve the regenerative potential of oaks. These fires encourage prolific sprouting of oak trees 

while limiting the growth of many oak competitors such as sugar maple (Acer saccharum), red maple 

(Acer rubrum), American beech (Fagus grandifolia), yellow-poplar (Liriodendron tulipifera), and 

black cherry (Prunus serotina). In addition, oak seedlings develop slowly in the low light of a closed 

overstory and thick understory. Increased light improves growth of oak species but it also encourages 

the growth of competing vegetation. Prescribed fire has been viewed as an effective tool to limit 

competing vegetation and improve the growth and form of oak regeneration in thinned versus 

unburned areas. In a study in the Shawnee Hills region of southern Illinois in the spring of 2003, the 

benefits of prescribed burning, mechanical thinning, and a combination of the two were examined. 

This poster includes the results of this study and implications for current and future silvicultural 

recommendations.

1Research Assistant, Southern Illinois University, Department of Forestry; 

2Associate Professor of Forestry, Southern Illinois University, Department 

of Forestry †Corresponding Author 1448 Makanda Rd, Makanda, IL 

62958: 618-319-0217, email: scottholevoet@yahoo.com.
2	 Fire 80 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Thinning and burning treatments influence within-canopy

leaf traits in seven hardwood species in Ohio

Jyh-Min Chiang1† and Kim J. Brown2

Leaf nitrogen content (LN, %) and leaf mass per area (LMA, g m-2) are important features that are 

closely linked to the photosynthetic performance of plants. Hence, models of ecosystem carbon 

balance often incorporate both leaf traits in simulations of potential net primary production. Forest 

management practices such as burning and thinning may change stand structure and soil dynamics, 

resulting in the changes in LN and LMA. For simulations at a larger spatial and temporal scale, 

additional information on species specific leaf traits and their plasticity to changing environments 

is needed. Our goal is to ultimately model the net effect of carbon sequestration under the future 

species rearrangement. The objective of this study was to understand how LN and LMA of different 

canopy tree species (Quercus alba L., Q. coccinea Muenchh., Q. prinus L., Q. velutina Lam., Carya

spp., Acer rubrum L., Liriodendron tulipifera L.) responded to thinning and/or burning treatments; 

and different landscape and soil properties in southern Ohio. We selected five trees for each of the 

seven species at each of the 12 experimental units (three replicates of control, thin, thin + burn, and 

burn treatments) of the Fire and Fire Surrogate (FFS) study. Leaves from the top and bottom of the 

crown profile were collected (N = 850) in the summer of 2003. We found significant effects (positive) 

of FFS treatments on LMA for most species at the bottom of the tree crown. FFS treatments also 

significantly increased LN of Q. coccinea and L. tulipifera at the bottom of the tree crown. Sun leaves 

showed consistent LN and LMA regardless of FFS treatments. Regression tree analysis showed no 

effects of landscape features and soil properties on LN and LMA. Interspecies differences accounted 

for most variations of LN and SLM. Our study suggested FFS treatments increased the net primary 

production of leaves at the lower crown position. Different species tended to retain their respective 

LN and SLM under different landscape and soil features. This finding will facilitate our efforts to 

model the impacts of future species redistribution of ecosystem carbon balances.

1

Ph.D. Candidate, Department of Environmental and Plant Biology, 

Ohio University; 2Assistant Professor, Department of Environmental 

and Plant Biology, Ohio University; †Corresponding author, 317 Porter 

Hall, Ohio University, Athens, OH 45701, 740-593-0405, email: 

jc293302@ohio.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 281

ESTIMATING FIRE BEHAVIOR FROM OVERHEAD IMAGERY IN EASTERN 

OAK FORESTS

Robert Kremens1†, Anthony Bova

2

, Matthew Dickinson3, Jason Faulring4, Shari McNamara5

We have used multiple-frame overhead infrared imagery to measure heat flux from a prescribed fire 

in Southeastern Ohio. These data were obtained using a specially designed wildfire airborne mapping 

camera that produces accurate geographic maps of short, mid, and longwave infrared emission along 

with very high-resolution visible images. These images may be overlaid to produce a total heat map. 

In-fire measurements were performed simultaneously with overhead image capture to provide a crosscalibration of ground leaving energy flux with sensor-reaching energy flux. This technique removes 

the effects of atmospheric absorption and uncertainties in instrument calibration. We correlated the 

heat release as measured by the airborne camera system to conventional measures of fire severity such 

as fuel burn-up. Forty plots were measured using conventional methods. Excellent agreement was 

obtained between conventional plot measurements of fuel consumption and the airborne heat-flux 

measurements. The airborne technique allows simultaneous observation of entire burn units and may 

provide a substantially better (higher accuracy, more synoptic view and lower cost) measure of fire 

characteristics than ground-based methods that are based on conventional sampling.

1

Senior Research Scientist, Rochester Institute of Technology; 2Physical 

Scientist, USDA Forest Service, Northern Research Station; 3Ecologist, 

USDA Forest Service, Northern Research Station; 4Integration Engineer, 

Rochester Institute of Technology; 5Graduate Student, Rochester Institute 

of Technology; †Corresponding author, 54 Lomb Memorial Drive, 

Rochester, NY 14623, 585-475-7286, email: kremens@cis.rit.edu.
2	 Fire 82 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

EFFECTS OF SILVICULTURAL TREATMENTS ON ACORN WEEVIL 

OCCURRENCE IN MIXED-OAK FORESTS

Jeffrey A. Lombardo1† and Brian C. McCarthy2

Oak regeneration failure in the hardwood forests of the Central and Eastern United States has been 

well documented. Strict policies of fire suppression likely have contributed to this decline by allowing 

shade-tolerant species to dominate the understory, suppressing the growth of oak seedlings and their 

subsequent release to larger size categories. What is not clear is the role that fire suppression may 

have played on the major oak seed predator, the acorn weevil (Coleoptera: Curculionidae). Weevils 

have been known to depredate a large percentage of acorns produced in a given growing season. 

This can result in lower seedling recruitment. The resulting low seedling density reduces the efficacy 

of the shelterwood-burn technique in regenerating oak stands. The purpose of this research was to 

determine the effects of three commonly used silvicultural treatments on weevil occurrence in mixedoak forests of southeastern Ohio. For this study we used the silvicultural treatments of the National 

Fire and Fire Surrogate study at the Zaleski State Forest and Vinton Furnace Experimental Forest 

sites, both of which are located in Vinton County, Ohio. The treatments include: prescribed fire, 

thinning, thinning followed by prescribed fire, and an untreated control. We tested the hypothesis 

that burning in the spring reduces the number of weevils in an area by inducing mortality on the 

adults as they emerged from the soil, and that thinning increases weevil activity by allowing more 

sunlight to penetrate the canopy, thus increasing ambient temperature. Adult weevils were captured 

using insect traps; acorns traps were used to determine the presence of larvae in acorns. Data from 

the insect traps support our hypothesis at the Vinton Furnace site; however, the Zaleski site had a 

more even distribution of weevil occurrence, signifying a possible site-by-treatment interactive effect. 

Results of this study show that fire suppression may negatively affect oak regeneration by allowing a 

greater occurrence of acorn weevils in a stand, thus reducing the number of new oak seedlings.

1

Graduate Student, Department of Environmental and Plant Biology, 

Ohio University; 2Professor of Forest Ecology, Department of 

Environmental and Plant Biology, Ohio University; †Corresponding 

author, 317 Porter Hall, Ohio University, Athens, OH 45701, 

740-597-3116, email: jl292804@ohio.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 283

Oak and maple stump sprout dynamics in response to thinning

and burning treatments in southern Ohio

Robert P. Long1†, David K. Apsley2, and Joanne Rebbeck1

Three southern Ohio sites (REMA, Tar Hollow, and Zaleski) are the focus for testing the efficacy 

of thinning (T) and the combination of thinning and burning (TB) on oak regeneration. In T and 

TB stands, sprouts on stumps (≥ 10 cm diameter) are an important component of regeneration 

dynamics. Thinning treatments, which reduced basal area by about 30 percent, were completed in 

winter-spring of 2000-2001. Stands were burned in March-April 2001. During 2004, we evaluated 

the density and growth of stump sprouts in sixteen 400-m2plots in each of six 20-ha treatment units. 

Treatment effects on red oak group (ROG = Quercus rubra, Q. coccinea, Q. velutina; n = 40 stumps) 

and white oak group (WOG = Q. alba, Q. prinus; n = 144 stumps) stump sprout density and growth 

were compared with their major competitor, red maple (Acer rubrum; n = 121 stumps). Analysis of 

variance showed that burning reduced the mean number of sprouts per hectare for all oaks and red 

maple combined. For instance, sprout densities were significantly (P < 0.006) lower in TB stands 

(491 sprouts/ha) than in T stands (875 sprouts/ha). There were consistently more red maple sprouts, 

which averaged 458/ha, across all six sites, compared with 62/ha for ROG species, and 162/ha for 

WOG species. However, these results may be related to the greater sprouting potential for smaller 

diameter red maple stumps, which averaged 25.5 cm (range: 10 to 50 cm) compared with mean 

stump diameters of 43.1 cm (range: 11 tot 73 cm) and 37.6 cm (range: 15 to 90 cm) for ROG and 

WOG stumps, respectively. Mean height and diameter growth for red maple, ROG, and WOG 

sprouts, evaluated by species group, was not significantly (P > 0.50) affected by treatments. The mean 

height of red maple sprouts, 3.3 m, was significantly (P < 0.001) greater than that of ROG and WOG 

sprouts, 2.5 and 2.0 m, respectively. Similarly, the mean basal diameter of red maple sprouts, 4.1 cm, 

was significantly greater than that of ROG and WOG sprouts, 2.2 and 2.0 cm, respectively. These 

results suggest that oak sprouts were not as competitive as red maple sprouts in thinned treatments, 

and that prescribed burning immediately after cutting did not improve the competitive status of oak 

stump sprouts.

1

USDA Forest Service, Northern Research Station; 2Ohio State University 

Extension; †Corresponding author, 359 Main Road, Delaware, OH 

43015-8640, 740-368-0050, email: rlong@fs.fed.us.
2	 Fire 84 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

IMPACTS OF REPEATED PRESCRIBED FIRES ON FUELS IN A CENTRAL 

APPALACHIAN HARDWOOD FOREST

Jessi E. Lyons1, Elizabeth Loucks2, Mary A. Arthur3†, David Loftis4

Throughout the central hardwood forest region, managers are increasingly using fire to attain 

various management objectives, including the promotion of oak regeneration, forest thinning, 

and fuel reduction. In the Daniel Boone National Forest in Kentucky, managers have used fire as a 

management tool in upland oak forests for more than 10 years. Funding for prescribed burning often 

comes with the justification that burning will reduce fuels and help prevent future unplanned fires. 

We examined fuel changes following one and two prescribed fires in spring. Fuel mass was sampled 

before the introduction of fire, after a single prescribed fire, after leaffall following one fire, and after 

two prescribed fires. Concerns about the potential for increased soil erosion following repeated fire 

prompted sampling of mineral soil exposure after the second prescribed fire. This study demonstrated 

that one and two prescribed fires decreased leaf litter, which was replaced in mass but not fuel 

continuity during leaffall. A single prescribed fire significantly decreased 10-h woody fuels, which 

were replenished the next leaffall in concurrence with leaf litter. Repeated prescribed fire significantly 

increased 1-h woody fuels. The duff layer decreased on burned treatments following a single fire, and 

decreased further after a second fire. After one burn, duff depth decreased more on subxeric landscape 

positions than on submesic positions. Mineral soil exposure also increased with number of fires. It 

was highest on subxeric and intermediate plots after one burn and higher on submesic plots after two 

burns. Subxeric plots showed a strong correlation between mineral soil exposure and litter consumed. 

Fuel consumption, duff depth, and mineral soil exposure varied with landscape position. On sites 

with steep slopes, repeated fire could lead to increased erosion due to exposed mineral soil. Common 

encounters with turkeys and their scratching has stimulated questions about their possible affinity for 

recently burned areas as a contributing cause for increased soil exposure.

1

Lab Technician, Department of Forestry, University of Kentucky; 2Former 

Graduate Student, Department of Forestry, University of Kentucky; 3

Associate Professor, Department of Forestry, University of Kentucky; 4

Research Forester, USDA Forest Service, Bent Creek Experimental 

Station; †Corresponding author, T.P. Cooper Building, Lexington, KY 

40546-0073, 859-257-2752, email: marthur@uky.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 285

FACTORS INFLUENCING WILDFIRE OCCURRENCE AND DISTRIBUTION 

IN EASTERN KENTUCKY

John K. Maingi1† and Mary C. Henry2

Most wildfires fires in Kentucky occur in the heavily forested Appalachian counties in the eastern 

portion of the state. These fires have been attributed to causes ranging from weather conditions to 

socioeconomic factors. We reconstructed a brief fire history of eastern Kentucky using Landsat TM 

and ETM+ images acquired between 1985 and 2002. We then examined relationships between fire 

frequency and area burned, and abiotic and human factors. Abiotic factors included Palmer Drought 

Severity Index (PDSI; negative values indicate drier conditions), slope, aspect, and elevation, and 

human factors such as county unemployment rates and distance to roads and populated areas. About 

83 percent of the total area burned experienced fire only once, 14 percent twice, and 3 percent 

three times. More fires burned in the winter (January-March) fire season than in the fall (OctoberDecember) season, but the latter fires were larger on average and accounted for about 71 percent of 

the total area burned. Fire size was negatively correlated with the mean PDSI in both fire seasons. 

There was a higher frequency of burning both at higher elevations and on steeper slopes. There 

was no relationship between aspect and frequency of burning. Despite the perceived link between 

jobless claims in the region and frequency of fires, we found no correlations between monthly 

unemployment rates and arson-caused fires. The number of fires recorded was negatively correlated 

with both distance of fire to road (p < 0.001) and distance to populated area (p < 0.001).

1,2

Assistant Professors, Department of Geography, Miami University; †

Corresponding author, 210 Shideler Hall, Oxford, OH 45056 USA, 

(513) 529-5024, email: maingijk@muohio.edu.
2	 Fire 86 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

HARDWOOD REGENERATION FOLLOWING PRESCRIBED FIRE AND 

THINNING IN MIXED-OAK FORESTS OF SOUTHEASTERN OHIO

Brian C. McCarthy1† and Matthew A. Albrecht2

Stand thinning and prescribed fire are widely used forest management tools throughout the 

hardwood forests of the Central Appalachians. Recent changes in historical disturbance regimes 

are hypothesized to have altered the structure and composition of many oak-dominated forests. In 

particular, there are few oaks in the regeneration layer (especially 1 to 4 inches d.b.h.) throughout 

the understory of most oak-dominated forests. Restoration efforts likely will require significant 

changes in disturbance regimes to alter the current successional trajectory. We examined the patterns 

of hardwood regeneration in a replicated field experiment over 5 years. Three forests were selected 

for study in southern Ohio. Within each forest, four adjacent 20 to 30-ha stands were selected, with 

one each allocated randomly to the following treatments: control, prescribed burn, thin, and thin 

followed by prescribed burn. Seedlings and saplings were sampled in replicated study plots in the year 

prior to treatment (2000), the year following treatment (2001), and 3 years later (2004). Structural 

data (density) were analyzed by analysis of covariance; compositional data (relative abundance) were 

analyzed by ordination procedures. Treatments had various effects dictated by species life history, 

reproductive episodes, and stochastic events. A single prescribed fire reduced seedling and sapling 

densities of red maple, the understory dominant in these forests. However, red maple rapidly 

recovered to pretreatment levels in all units four growing seasons after the disturbances. Mechanical 

thinning treatments accelerated understory recruitment of early successional, shade-intolerant tree 

species that regenerated from seed, e.g., tuliptree, and resprouted from a seedling bank, e.g., sassafras. 

Tuliptree also was strongly influenced by burning, resulting in greatly enhanced reproduction and 

advancement into the sapling size class via fast growth rates. White oak showed no response, black 

oak increased slightly, and chestnut oak varied moderately with respect to treatment. Overall, the 

treatments were effective at advancing a greater proportion of tuliptree into the succeeding forest. 

However, the effects of treatment on oak regeneration are unclear, so additional longitudinal data will 

be needed. Subsequent periodic fires may help to slow mesic competitors over the long run.

1

Professor of Forest Ecology, Department of Environmental and 

Plant Biology, Ohio University, 2Graduate Student, Department of 

Environmental and Plant Biology, Ohio University; †Corresponding 

author, 416 Porter Hall, Ohio University, Athens, OH 45701, 

740-593-1615, email: mccarthy@ohio.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 287

SOIL RESPIRATION RESPONSES TO TEMPORAL, TOPOGRAPHIC, AND 

SILVICULTURAL FACTORS IN A SOUTHEASTERN OHIO FOREST

Dawn R. McCarthy1† and Kim J. Brown2

Soil respiration (CO2 efflux) is an important component of carbon loss from forest ecosystems. 

Attempts to understand the relationship between silvicultural practices and soil respiration are 

complicated by physical and biological heterogeneity at the landscape scale. We examined the effects 

of burning on soil temperature, soil moisture, and respiration while controlling for slope position 

and thinning intensity. Treatments included unburned control, low intensity fire, high intensity 

fire, lime fertilization, and leaf litter removal. These treatments (each 2×2 m) were replicated in 

20 experimental blocks at different slope positions (upper vs. lower) and thinning treatments 

(closed canopy vs. 50-percent removal) in a mixed-oak forest in southeastern Ohio. Monthly field 

measurements were conducted from April to November in 2004. Treatment and slope position were 

significant factors principally during the growing season. Upslope and thinned plots experienced 

midsummer soil-water reductions of 6 to 10 percent, respectively. Leaf litter removal reduced 

respiration by 15 percent compared to the control, while lime fertilization increased respiration by 10 

percent compared to the control. From the entire season’s data we developed a nonlinear model of soil 

respiration as a function of soil temperature for each treatment. Model equation parameters differed 

only for slope position and fertilization treatment, revealing greater sensitivity in respiration rates at 

lower slopes compared to upper slopes. Overall, thinning did not significantly affect soil respiration, 

but there was a trend toward reduced soil respiration rates (P = 0.09) in the low-intensity burn plots. 

Resulting values for soil moisture and soil temperature are a function of both topography and land 

use, leading to unequal effects of thinning and burning across the landscape. While the current study 

is informative with respect to the microclimatological mechanisms of thinning and burning on soil 

respiration, the small treatment area prevents a large-scale extrapolation to a thin/burn scenario due 

to other factors, e.g., mechanical soil disturbance.

1

Graduate Student, Department of Environmental & Plant Biology, Ohio 

University; 2Assistant Professor, Department of Environmental & Plant 

Biology, Ohio University; †Corresponding author, 317 Porter Hall, Ohio 

University, Athens, OH 45701, 740-707-1342, email: db642701@ohio.edu.
2	 Fire 88 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

CHANGES IN THE STAND STRUCTURE AND SPECIES COMPOSITION OF 

OAK-DOMINATED COMMUNITIES FOLLOWING A PRESCRIBED BURN

Virginia L. McDaniel†, Michael A. Jenkins, and Robert N. Klein

Fire suppression has affected many forest communities in Eastern North America. It has played 

a significant role in deterring regeneration of oaks and encouraging regeneration of more shadetolerant species in oak dominated communities. At Kings Mountain National Military Park in 

South Carolina, fire suppression has altered stand structure and species composition on oakdominated ridges. It has given shade-tolerant, fire-intolerant species a competitive advantage over 

shade-intolerant, fire-tolerant species. In 2000, we sampled 19 plots prior to a prescribed burn 

and resampled the same plots a year after burning. Prior to burning, the overstory was dominated 

by oaks, but advanced regeneration was dominated by blackgum, red maple, and sourwood. The 

pre-burn diameter distribution of oak species was bell-shaped, but the distribution of other species 

(including black gum, sourwood, and red maple) was reverse J-shaped. This indicates that understory 

composition in fire suppressed stands has shifted from dominance by relatively xeric oak species to 

dominance by more mesic species. One year after the burn, the oaks continued to have a bell-shaped 

distribution, but the number of small stems of fire-intolerant species was greatly reduced, resulting in 

a more bell-shaped distribution.

Twin Creeks Natural Resources Center, Great Smoky Mountains National 

Park; †Corresponding author, 1314 Cherokee Orchard Rd., Gatlinburg, 

TN 37738, 865-430-4751, email: virginia_mcdaniel@nps.gov.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 289

experimental evaluation of fire history reconstruction

using dendrochronology in white oak (Quercus alba)

Ryan W. McEwan1, Todd Hutchinson2, Robert Ford3, Brian C. McCarthy4†

Fire history often is reconstruction through dendroecological analysis of fire scars on tree boles. An 

understanding of historical fire regimes is critical for understanding current forest structure and forest 

dynamics, and for evaluating forest management practices. Even so, the validity of this approach has 

rarely been evaluated experimentally. Do we know whether fire scars accurately record fire history 

in dendroecological samples? We collected basal cross sections from stands where prescribed fires 

had occurred to examine the efficacy of fire history reconstruction in eastern hardwood forests. We 

collected samples from 82 trees in two sites, each having experienced three fire treatment intervals 

(frequent, infrequent, and control) over the last nine years. Prescribed fires were set in 15 area/time 

combinations. Of these, 10 were recorded as fire scars. In all areas, the first year of prescribed fire 

was recorded in at least one of the samples, and scarring occurred when there had been pauses in the 

burning regime of more than 3 years. Although the known fire history generally was reconstructed 

through tree-ring analysis, the proportion of samples bearing scars was exceedingly low (12 percent). 

Fires that occurred in subsequent years generally were not recorded in the tree-ring record (one of 

six cases), suggesting that multiyear fuel accumulation is necessary to create fire intensity needed to 

wound trees. Our study suggests that tree-ring analysis is an effective tool for reconstructing burning 

regimes when the fire-free interval is long enough to allow for fuel accumulation; however, a frequent 

burning regime (intervals of 1 to t3 years) may be undetectable.

1

Doctoral Candidate, Ohio University, Department of Environmental 

and Plant Biology; 2Research Ecologist, USDA Forest Service, 

Northeastern Research Station; 3Dendrochronologist, USDA Forest 

Service, Northeastern Research Station; 4Professor of Forest Ecology, 

Ohio University, Department of Environmental and Plant Biology; †

Corresponding author, 315 Porter Hall, Ohio University, Athens, OH 

45701, 740.593.1615, email: mccarthy@ohio.edu.
2	 Fire 90 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

IMPACT OF FIRE ON SOIL RESOURCE PATTERNS IN MIXED-CONIFER 

FORESTS IN THE SOUTHERN CASCADE RANGE OF NORTHERN 

CALIFORNIA

Jessica Miesel1†, Carl Skinner2, and Ralph Boerner3

The montane ecosystems of the Cascade Range have been subjected to repeated manipulation and 

active fire suppression for more than a century. This has resulted in changes in community structure 

that contribute to increased wildfire hazard and severity. Ecosystem restoration to reduce the wildfire 

hazard has received substantial attention in recent years, though many ecological questions remain 

unanswered. This study addresses below-ground impacts of restorative treatments. We report 

preliminary results of the application of prescribed fire and the combination of fire and thinning on 

soil chemical and microbial parameters in treatment units of 10 ha each in the Klamath National 

Forest of northern California. Soil pH and total inorganic N increased in burned units (with and 

without thinning) one year after treatment, but differed significantly between treatments. Soil organic 

C decreased and C:N ratio increased as a result of fire in the burn-only treatment; no significant 

changes were observed for thin+burn treatment plots. Nitrogen mineralization rates did not change as 

a result of fire for both burn-only and thin+burn plots. Activity of acid phosphatase was reduced by 

fire, with and without thinning, whereas activity of chitinase was reduced by fire in thin+burn plots 

only. Changes in levels of phenol oxidase activity as a result of fire were not significant for treatments 

with or without fire. Our study contributes to an understanding of the ecological effects of fire on soil 

chemical and microbial properties. This project is part of the interdisciplinary national Fire and Fire 

Surrogate Network study and will provide information for informed land management decisions.

1

Graduate Research Assistant, The Ohio State University, Department of 

Evolution, Ecology, and Organismal Biology; 2Geographer, USDA Forest 

Service Pacific Southwest Research Station; 3Professor, The Ohio State 

University, Department of Evolution, Ecology, and Organismal Biology; †

Corresponding author, 300 Aronoff Laboratory, 318 W. 12th Ave., 

Columbus, OH 43210, (614) 292-9373, email: miesel.1@osu.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 291

Avian community responses to prescribed burns and

shelterwood harvests

Donald B. Miles1†, Henry Streby2, and Dan Weber2

Previous forest management practices in U. S. Forests emphasized the suppression of fires. A 

consequence of this management practice has been the excessive accumulation of fuel and high tree 

densities. To avoid severe and catastrophic fires, resource managers now use a variety of treatments to 

diminish fuel loads. Alternative treatments include prescribed burns and shelterwood harvests, both 

of which are assumed to mimic natural ecosystem function. However, avian responses to silviculture 

treatments have been investigated only recently. We present the results of a study to determine 

whether avian community composition, productivity, and foraging behavior are affected by fire and 

fire surrogate treatments. Avian species composition and abundance in each of four treatment types 

were estimated using point-count census methods. The treatment plots were control, burn, thin, and 

burn + thin. The treatments were replicated at three sites. We also determined differences in nest 

productivity. Avian censuses were conducted 1 year before the treatments to obtain baseline data 

on species composition and relative abundance, and 3 years posttreatment. Our results show acute 

and relatively long-term responses to forest disturbance. We compared community composition, 

nest productivity, and foraging behavior using data from the baseline samples to determine whether 

species composition was homogeneous among the treatments. Species composition was significantly 

different among treatments, with the highest diversity in thin and burn plots. However, there was 

considerable heterogeneity in the data. Average nest productivity also varied across treatments. We 

document how species composition shifts in response to the thin and burn treatments.

1

Professor of Vertebrate Zoology, Ohio University, Department of 

Biological Sciences, †Corresponding author, Athens, Ohio 45701, 2

Graduate Research Biologist, Department of Biological Sciences, Ohio 

University, Athens, OH 45701, 740-593-2317, email: dmiles2@ohio.edu.
2	 Fire 92 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

REHABILITATION OF AN UPLAND HARDWOOD FOREST USING 

PRESCRIBED FIRE AND HERBICIDE

Rebecca Montgomery1†, Jeff Earl1, Chris Stuhlinger2, Matthew Pelkki3, and Kyle Cunningham4

Many upland hardwood forests have been degraded due to improper management (high-grading) or 

insect/disease outbreaks. The absence of periodic fires has changed the forest composition to a higher 

proportion of undesirable shade-tolerant species. Some desirable species such as oaks have declined. 

Methods are needed to rehabilitate these forests by encouraging the regeneration and growth of 

desirable tree species and controlling undesirable species. The application of prescribed fire and 

herbicide are being evaluated in an Ozark upland hardwood forest in northwestern Arkansas. In a 

landscape where the wildland-urban interface is expanding rapidly, concern is increasing with respect 

to the use of herbicides and fire as management tools. The advantages and limitations of each method 

must be understood so that appropriate recommendations can be made for specific situations. In 

this study, treatment plots were left alone, burned in the spring, treated with Chopper® herbicide in 

the fall to kill undesirable trees, or treated with both fire and herbicide. Fire intensity on the burned 

plots was measured with Tempilaq® (Tempil, South Plainfield, NJ) temperature-indicating liquids 

applied to metal tags, and varied widely depending on available fuels. Fire intensity (temperature) 

can be related to the understory kill of small-diameter stems and subsequent regeneration of 

desirable species. Early observations indicate successful control of larger stems by herbicide, while 

burning may only temporarily control sprouting and growth of small undesirable stems. Prescribing 

repeated burns, or applying herbicide, might achieve the desired result of restoring a significant oak 

component to these forests.

1

Program Technicians, University of Arkansas – Monticello, School of 

Forest Resources; 2University System Forest Manager, Arkansas Forest 

Resources Center; 3George H. Clippert Endowed Chair, University of 

Arkansas – Monticello, School of Forest Resources; 4Forestry Instructor, 

University of Arkansas, Cooperative Extension Service; †Corresponding 

author, P.O. Box 3468, Monticello, AR 71656, 870-460-1052, email: 

montgomeryr@uamont.edu. 
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 293

ALTERED DISTURBANCE REGIMES: DEMISE OF FIRE IN THE EASTERN 

UNITED STATES

Gregory J. Nowacki1† and Robert A. Carr2

A series of maps was generated to help alert and educate people to the pervasiveness of fire regime 

changes across the Eastern United States. Using geographic information systems, we assigned fire 

regimes to spatial vegetation databases to depict past and current conditions. Comparisons revealed 

substantial reductions in fire throughout the East. The most dramatic shifts took place in the former 

midwestern grasslands and across a broad swath of Southern and Central States, where pine and oak 

communities historically dominated. Land-use changes, e.g., agricultural and forest-type conversions, 

and 20th century fire suppression largely explain these shifts. Fire regime change was somewhat 

limited in northern climes (northern hardwood systems), in the mixed mesophytic region, and within 

the Mississippi Embayment. Unforeseen consequences of prolonged fire suppression are mounting 

while restoration opportunities are waning.

1

Regional Ecologist and 2Regional GIS Planning Analyst, USDA Forest 

Service, Eastern Region; †Corresponding author, 626 E. Wisconsin Ave., 

Milwaukee, WI 53202, 414-297-1977, email: gnowacki@fs.fed.us.
2	 Fire 94 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

LIGHTNING-INITIATED GROWING SEASON FIRES CHARACTERIZED OAKDOMINATED ECOSYSTEMS OF SOUTHERN OHIO 

Sheryl M. Petersen1 and Paul B. Drewa2†

The persistence of oak-dominated ecosystems throughout the Central Hardwoods Region has 

been attributed to chronic, noncatastrophic fire regimes prior to Euro-American settlement. The 

nature of these fires is not entirely understood. We used lightning strike, thunderstorm activity, and 

precipitation data to elucidate the seasonal timing of natural fire regimes for southern Ohio. Our 

results suggest that fires are most likely to be initiated during the late growing season (especially 

August) when there is a high frequency of lightning strikes and thunderstorm activity that occurs 

simultaneously with dry ground-cover fuel conditions. We hypothesize that the use of prescribed 

growing-season fires will effectively deter encroachment of shade-tolerant hardwood vegetation and 

foster oak regeneration. Such fires will provide an evolutionary basis for the conservation of oakdominated ecosystems of southern Ohio.

1

Ph.D. Graduate Student; 2Assistant Professor, Department of Biology, 

Case Western Reserve University; †Corresponding author, Department of 

Biology, Case Western Reserve University, 10900 Euclid Ave., Cleveland, 

OH 44106, 216-368-4288, email: paul.drewa@case.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 295

USE OF TEMPERATURE-SENSITIVE PAINTS AS AN INDEX OF HEAT 

OUTPUT FROM FIRE

Joanne Rebbeck1†, Daniel Yaussy2, Louis Iverson3, Todd Hutchinson3, Robert Long4, Anthony Bova

5

and Matthew Dickinson6

Early spring, dormant-season prescribed surface fires were conducted in 2001 and repeated in 2005 

within thinned and three unthinned mixed-oak forests of southeastern Ohio. In all, 120 ha were 

burned on three forests. Research has shown that maximum temperatures reached by measurement 

devices placed in fires can be useful in evaluating fire effects on vegetation. Most commonly used are 

tags painted with Tempilaq® temperature sensitive paints (Tempil Inc., South Plainfield, NJ). Paints 

melt and change color at specific temperatures. This low-cost ($0.05 per unit), “low tech” method 

of measuring the maximum temperature was compared with a high-cost ($110 per unit), high-tech 

Type K® thermocouple probes and Hobo® data logger (Onset Computer Corporation, Bourne, MA) 

system. Maximum temperatures have been shown to indicate fuel consumption and fireline intensity.

Aluminum tags (2.5 x 7 cm) were painted with six (2001) or nine (2005) Tempilaq paints ranging 

from 79° to 427°C. Tags were hung at the same height (25 cm) and location as the probe tips. 

Data loggers were programmed to record probe temperature every 1 to 2 seconds. There was very 

good agreement between the maximum temperatures recorded by the two devices during both fires. 

Maximum temperatures were higher in the 2005 than in the 2001 fires (the first fires on these sites 

in decades). During 2001, maximum temperatures in the burn only units were higher than those in 

the thined and burned units. No treatment differences were observed in 2005. The impacts of more 

intense fires on vegetation are being monitored. Paint tags indicate maximum temperatures and are 

inexpensive, but thermocouple probes provide more information and better correlations with fire 

behavior. 

1

Plant Physiologist, 2Forester, 3Ecologist, 4Plant Pathologist, 5Physical 

Scientist, and 6Ecologist, USDA Forest Service, Northern Research 

Station; †Corresponding author, 359 Main Road, Delaware, OH, 43015, 

740-368-0054, email: jrebbeck@fs.fed.us
2	 Fire 96 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

IMPACT OF FIRE, SILVICULTURAL MANIPULATION, AND MICROSITE

CONDITIONS ON WOODY DEBRIS DECAY DYNAMICS IN 

A MIXED-OAK FOREST

Darrin L. Rubino1†, Lindsey E. Hesch2, and Brian C. McCarthy3

Woody debris (downed limbs and boles) is an important component of forested ecosystems. It 

serves as habitat for a wide variety of organisms and plays an important role in nutrient cycling and 

dynamics. However, little is known about the wood-decay process, especially in the topographically 

complex mixed-oak forests of Eastern North America. The goal of this investigation was to determine 

how fire and silvicultural management affect the decay rate of woody debris by using wood blocks 

cut from commercially available native lumber. This study was conducted in Zaleski State Forest 

in southeastern Ohio (an Ohio Hills study site in the long-term Fire and Fire Surrogate study). 

Following silvicultural treatment and burning, wood blocks of “red” oak (Quercus L. subgenus 

Erythrobalanus) and “white” oak (Quercus L. subgenus Lepidobalanus) were placed on the forest floor 

in burned, thinned, thinned + burned, and control (no manipulation) units. In each unit, blocks 

were placed in xeric, e.g., ridges and southwest-facing slopes, and mesic, e.g., valleys and northeastfacing slopes, microsites to also investigate the impact of moisture availability on the decay process. 

Site moisture availability was determined using the integrated moisture index. After 27 months of 

decay, mean mass loss was significantly (P < 0.05) greater for red than white oak. Mean mass loss was 

51.9 ± 1.7 (SE) percent for red oak and 36.3 ± 1.3 percent for white oak. Using a two-way ANOVA 

to compare the mean percent mass loss in each unit, microsite, and microsite × unit combination 

revealed no significant differences in decay rate for white oak. Alone, microsite and unit had no 

significant affect on decay rate in red oak. However, the microsite × unit interaction was observed 

for red oak. The fastest decay rates were observed in mesic microsites in the thin + burn unit and the 

slowest in the mesic microsites in the thinned unit. Decay dynamics appear to be species specific and 

fire and silvicultural manipulation may lead to complex changes in decay dynamics in topographically 

dissected mixed-oak forests of Eastern North America. 

1

Assitant Professor of Biology, Biology Department, Hanover College; 2

Graduate Assistant, Botany Department, Miami University of Ohio; 3

Professor of Forest Ecology, Department of Environmental and Plant 

Biology, Ohio University; †Corresponding author, P.O. Box 890, Hanover, 

IN 47243-0890, 812-866-7247, email: rubino@hanover.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 297

EFFECTS OF FOREST MANAGEMENT ON SOIL SEEDBANK SPECIES 

COMPOSITION IN a MIXED-OAK FOREST in SOUTHEASTERN OHIO

Lisa R. Schelling1† and Brian C. McCarthy2

Burning and thinning are common management practices in mixed-oak forests. The impacts of these 

practices on the soil seedbank have not been examined. This information is important to consider to 

evaluate the full impact of these treatments on plant community dynamics in the understory. Species 

composition of the soil seedbank was examined under four forest management treatments: thin, 

burn, thin followed by burning, and an untreated control. Sampling areas were located in Vinton 

County, Ohio, at Zaleski State Forest and the Raccoon Ecological Management Area. Both sites are 

part of the USDA’s Fire and Fire Surrogate study. Thinning was conducted in the fall of 2000 and 

burns were conducted in the spring of 2001. Soil samples were collected in March 2004, 3 years 

following treatment. Samples were placed in the greenhouse and the emergence method was used to 

assess the species composition of seeds present in the soil. Seventy-one species were found in the soil 

seedbank. Ruderal and early successional species such as fireweed (Erechtites hieraciifolia (L.) Raf.), 

sedges (Carex spp.), and brambles (Rubus spp.) were the most commonly occurring species across all 

treatments. Multivariate analysis techniques revealed no grouping by stand management treatment. 

Log-linear analysis showed a treatment by functional group effect which, while not statistically 

significant (P = 0.0872), suggests possible biological significance. In different treatments there were 

different proportions of the various functional groups (grasses, perennial herbs, etc.). There seems to 

be no detectable difference in the species composition of the soil seedbank after short-term exposure 

to different forest management treatments, probably because the species that maintain a seedbank in 

the soil tend to be early successional. These species are not particularly sensitive to disturbance, but 

usually respond positively to disturbance. More disturbance-sensitive forest herb species tend not to 

maintain a soil seedbank, and thus did not register a response in this study. Studies examining aboveground species dynamics likely are sufficient to assess the impact of forest management on understory 

dynamics.

1

Graduate Student, Department of Environmental and Plant Biology, 

Ohio University; 2Professor of Forest Ecology, Department of 

Environmental and Plant Biology, Ohio University; †Corresponding 

author, 315 Porter Hall, Ohio University, Athens, OH 45701, 

740-597-3116, email: ls903103@ohio.edu.
2	 Fire 98 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

OAK-GRASSLAND RESTORATION DEMONSTRATION AREA PROJECT

Leslie M. Smith1† and James R. McCoy2

Land Between the Lakes is home to two oak-grassland restoration demonstration areas. The 

demonstration areas create conditions over a large contiguous landscape to demonstrate the feasibility 

of ecological restoration of an oak-grassland forest and the benefits it can provide to native wildlife 

and public recreation. Historical accounts and ecological research indicates these areas had an open 

oak canopy and understory dominated by grasses created and maintained through the use fire by the 

American Indians. The demonstration area totals about 8,000 acres divided into the north unit in 

Kentucky (3,000 acres) and the south unit in Tennessee (about 5,000 acres). The units are located 

near educational outreach facilities that will encourage public interest and learning opportunities. 

Treatments will focus on the tree thinning and prescribed fire. Fire effects monitoring and research 

will continue throughout the restoration effort. “Core” areas adjacent to the demonstration areas serve 

as controls to gauge the effectiveness of the restoration. Nearly 350 acres in the southern restoration 

unit were treated with prescribed fire in the spring of 2005, and 70 acres of timber have been marked 

for thinning. Fire-effects monitoring plots have been established and observations are continuing. 

1

Wildlife Biologist-Trainee, USDA Forest Service, Land Between the 

Lakes; 2Wildlife Biologist/Fire Management Officer, USDA Forest Service, 

Land Between the Lakes; †Corresponding author, 100 Van Morgan Drive, 

Golden Pond, KY 42211, 270-924-2084, email: lesliemsmith@fs.fed.us.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 299

INTEGRATED UPLAND HARDWOOD RESEARCH IN ARKANSAS’ 

BOSTON MOUNTAINS

Martin A. Spetich1

A series of interrelated studies is examining upland hardwood forest dynamics and development 

in the southwestern lobe of the Central Hardwood Region. The overall objective of this research 

is: 1) address practical, applied forest management needs based on a combination of current 

scientific thought and needs expressed by field practitioners, and 2) disseminate results to scientists, 

practitioners, and the public in ways that will benefit both society and the environment. The studies 

are addressing prescribed fire and fire history, growth, woody species reproduction, competitive 

capacity, stand dynamics, stand composition, forest species restoration, quantitative silviculture, 

development of forest management methods, forest ecology, disturbance ecology, and diversity of 

these upland hardwood forests.

1

USDA Forest Service, Southern Research Station, Arkansas Forestry 

Sciences Laboratory, P.O. Box 1270, Hot Springs, AR 71902, 

501-623-1180 Ext. 105, email: mspetich@fs.fed.us.
300 	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

ASSIGNING FIRE REGIMES ON THE MONONGAHELA NATIONAL FOREST, 

WEST VIRGINIA

Melissa Thomas-Van Gundy1

The use of prescribed fire on the Monongahela National Forest is expected to increase as the role of 

fire is better understood and appreciated. Ecological knowledge of an area’s fire regime is essential 

in meeting the intent of a burn program. A fire-adapted vegetation model emulating the coarsescale work was developed in 2002 as a first attempt to assign fire regimes on the Monongahela. 

Spatial analyses and maps were generated using geographic information systems, specifically the 

Model Builder extension in ArcView 3.2 (ESRI, Redlands, CA). Available resource themes were 

reviewed for relevancy in estimating fire regimes. From these, four themes were selected: land type 

associations, fire-adapted vegetation, potential natural vegetation, and current forest types. All themes 

were converted to 20- by 20-m grids for applicability at the stand level (10’s to 100’s of ha). Selected 

features in the themes were given scaling values of 1 through 5 to represent vegetation fire adaptation, 

with 1 representing vegetation most adapted to fire and 5 the least. Weights were assigned to each 

theme according to its estimated influence or importance on fire regimes. Current forest type and 

potential natural vegetation were weighted equally and higher than other input themes. The resulting 

weighted scores were then categorized into fire regime groups based on ecological knowledge of fire-

vegetation relationships. Fire regimes were assigned as follows: model ranking one = fire regime IV 

(35 to 100 years, stand-replacement severity), model ranking two = fire regime I (0 to 35 years, low 

severity), model ranking three = fire regime III (35 to 100 years, mixed severity), and rankings four 

and five = fire regime V (200+ years, stand-replacement severity). Fire regime II (0 to 35 years, standreplacement severity) was not considered applicable to landscapes on the Monongahela. 

1

Forest Ecologist, Monongahela National Forest, 200 Sycamore Street, 

Elkins, WV 26241, 304-636-1800, email: mthomasvangundy@fs.fed.us.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 301

INFLUENCE OF PRESCRIBED FIRE ON STEM GIRDLING AND MORTALITY

Jeffrey S. Ward1† and Patrick H. Brose2

This study examined the response of individual stems to prescribed fire in the eastern oak (Quercus 

spp.) forest. Following seven prescribed fires in Connecticut, all stems at least 4.5 feet tall were tallied 

within thirty 45- by 45-foot plots. To increase the sample of stems with diameters of at least 2.5 

inches, trees that were within 45 feet of each plot center but outside of the interior plots, also were 

measured. Diameters were measured to the nearest 0.04 inches for all stems with diameters of at least 

0.4 inch. Heights were measured to the nearest 0.4 inch to a maximum of 80 inches. The fraction of 

each stem girdled by fire was recorded in increments of 10 percent. Trees with dead tops (topkilled) 

were recorded as 100-percent girdled. In all, 3,476 stems were examined. Temperatures were recorded 

during the fires with thermocouples at each plot corner and at plot center. Average maximum 

temperatures within plots was 360oF (range: 142 to 696oF). Average stem girdling was 95 percent 

or more for seedlings (< 0.4 inch diameter) regardless of species. Average stem girdling decreased by 

approximately 5 percent per 1 inch increase in diameter for larger stems. Less girdling was noted for 

oaks than for other species in the poletimber size class (4.5 to 10.5 inches diameter). Some sawtimber 

trees (> 10.5 inches diameter) also were injured by prescribed fire. Twenty percent of sawtimber oaks 

had at least 25-percent girdling compared with 28 percent for non-oak sawtimber. A similar pattern 

was noted for stems that were topkilled. There was no difference among species in the proportion of 

saplings (0.4 to 4.5 inches diameter) that were topkilled. Fewer poletimber and sawtimber oaks were 

topkilled than non-oak species. Topkilled oaks sprouted more frequently than non-oaks, 68 vs. 44 

percent, respectively. However, nonoak sprouts were more numerous and taller. For all species, both 

girdling and topkill increased as the average maximum temperature of the prescribed fire increased. 

For example, 45 percent of trees in the 4-inch diameter class were topkilled when exposed to 

temperatures > 300oF compared with only 5 percent for stems exposed to temperatures < 300oF. 

1

Chief Scientist, The Connecticut Agricultural Experiment Station, 

Department of Forestry and Horticulture; 2Forester, USDA Forest 

Service, Northern Research Station; †Corresponding author, PO Box 

1106, New Haven, CT 06504, 203-974-8495, email: 

jeffrey.ward@po.state.ct.us.
30	 Fire 2 in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1

Putting Fire on the Ground with Prescribed Burning

Associations

John R. Weir1†, Terrence G. Bidwell2, and Chris M. O’Meilia3

A prescribed burn association is a group of landowners and other concerned citizens that form a 

partnership to conduct prescribed burns. Prescribed burning is the key land management tool used to 

restore and maintain native plant communities to their former diversity and productivity for livestock 

production and wildlife habitat. There are four excuses that are used by land managers when asked 

why they do not use prescribed fire: liability, no training or experience, not enough labor, and not 

enough equipment. Prescribed burning associations can be the answer to all of these excuses. To start 

an association, get a group of interested people together. Be sure to involve key landholders, agency 

personnel, and fire departments within the community. Associations already have been formed that 

have developed goals and guidelines that other groups can use. The association should incorporate 

so that all dues, donations, and gifts are tax deductible and the association is eligible for grants from 

public and private groups. Some local rural fire departments assist the associations during burning 

operations. This cooperative effort provides the association with added equipment and personnel 

and provides the fire department with training time, possible added income, and community service. 

The primary benefit of a prescribed burning association is its ability to influence politics. Also, the 

increased use of prescribed fire will influence the continued safe use of fire with future generations. At 

this time there are eight prescribed burning associations conducting fires in Oklahoma.

1

Superientendent, OSU Research Range, OAES Field and Research 

Services Unit; 2Professor and Extension Specialist, Rangeland Ecology 

and Management, OSU; 3Biologist, US Fish & Wildlife Service; 

†

Corresponding author, 4922 South Coyle Road, Stillwater, OK 74074, 

405-743-4714, email weir@okstate.edu.
Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1 303

FIRE RESEARCH ON THE RACCOON ECOLOGICAL MANAGEMENT AREA

Daniel Yaussy1†, Todd Hutchinson2, Matthew Dickinson2, Joanne Rebbeck3

The Raccoon Ecological Management Area (REMA) consists of 16,000 acres of forest located on 

the unglaciated Allegheny Plateau in southeastern Ohio. The area was cutover to provide charcoal 

to the iron furnaces that operated during the mid 19th century. The overstory is dominated by oaks 

with abundant red maple in the midstory and understory. Research on the use of fire to restore 

a mixed-oak ecosystem was initiated on the REMA in 1995 with a grant from the USDA Forest 

Service’s Ecosystem Management Program. Periodic and annual prescribed burns were compared with 

respect to their effect on flora and fauna. These low-intensity fires did not increase the light reaching 

the forest floor sufficiently to promote oak regeneration. The Joint Fire Science Program provided 

funding for a replication of the national Fire and Fire Surrogate study, which was installed on the 

REMA in 2001 as part of the Ohio Hills site. This ongoing study combines thinning from below 

and prescribed fire to provide varying light levels and oak competition control to promote mixed-oak 

ecosystem sustainability. Many additional investigations by Forest Service and university scientists 

have been added to the original experimental design, which increases insight into the effects of these 

management practices. The processes determining fire behavior and effects are being modeled with 

funds provided by the National Fire Plan. Fire monitoring technologies and fire effects models are 

being developed. A hydrology and ecosystem process model is being applied to the sites to provide 

fuel characteristics for fire modeling. A study is being established on the REMA in which herbicide 

will be used prior to harvesting under a shelterwood system to control the sprouting of the polesized oak competitors. Prescribed fire will be used to control new seedling competition 3 to 5 years 

following a major acorn crop. It is hypothesized that the combination of these management practices 

will promote sustainability of the mixed-oak ecosystem. The research of fire effects and behavior on 

the REMA have made it on of the premier sites for producing information on the management of 

mixed-oak ecosystems in the United States.

1

Forester, 2Ecologists, and 3Plant Physiologist, USDA Forest Service, 

Northern Research Station, †Corresponding author; 359 Main Road, 

Delaware, OH 43015, 740-368-0101, email: dyaussy@fs.fed.us.
Printed on Recycled Paper

Dickinson, Matthew B., ed. 2006. Fire in eastern oak forests: delivering 

science to land managers, proceedings of a conference; 2005 November 

15-17; Columbus, OH. Gen. Tech. Rep. NRS-P-1. Newtown Square, PA: U.S. 

Department of Agriculture, Forest Service, Northern Research Station. 303 p.

Contains 20 papers and 36 poster abstracts presented at a conference on fire in 

oak forests of the Eastern United States that was held at the Ohio State University, 

Columbus, Ohio, on November 15-17, 2005.
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(Not all prohibited bases apply to all programs.) Persons with disabilities who require alternate means 

for communication of program information (Braille, large print, audiotape, etc.) should contact USDA’s 

TARGET Center at (202) 720-2600 (voice and TDD). To file a complaint of discrimination, write to 

USDA, Director, Office of Civil Rights, 1400 Independence Avenue, S.W., Washington, DC 20250-

9410, or call (800)795-3272 (voice) or (202)720-6382 (TDD). USDA is an equal opportunity provider 

and employer. 

Capitalizing on the strengths of existing science capacity in the 

Northeast and Midwest to attain a more integrated cohesive 

landscape scale research program