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 0.5to
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 μ2
Ch: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
10	 Fire 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
DEPTHINSEDIMENT(cm)
ca. 1700 AD
650 ± 60
4210 ± 40
C-14AGE(BP)
20
%
0
PIN
E
20 40 60
%
0
O
A
K
%
0
G
R
A
SSES
20
%
0
A
G
R
IC
ULTUR
AL
IN
DICA
TO
RS
200 400
microns^2 : upland pollen
0
C
HA
RC
O
A
L
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
11Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
DEPTHINSEDIMENT(cm)
ca. 1750 AD
730 ± 60
C-14AGE(BP)
20 40 60
%
0
PIN
E
20 40
%
0
O
A
K
%
0
G
R
A
SSES
20
%
0
A
G
R
IC
ULTUR
AL
IN
DICA
TO
RS
200 400 600 800
microns^2 : pollen
0
C
HA
RC
O
A
L
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
DEPTHINSEDIMENT(mm)
ca. 1680 AD
2180 ± 130
C-14AGE(BP)
20
%
0
PIN
E
20 40 60
%
0
O
A
K
%
0
G
R
A
SSES
20
%
0
A
G
R
IC
ULTUR
AL
IN
DICA
TO
RS
1000 2000
microns^2 : upland pollen
0
C
HA
RC
O
A
L
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
13Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
DEPTHINSEDIMENT(cm)
ca. 1800 AD
20 40
%
0
W
H
ITE
PIN
E
20
%
0
PITCH
PINE
20 40
%
0
O
A
K
%
0
G
R
A
SSES
20
%
0
A
G
R
IC
ULTUR
AL
IN
DICA
TO
RS
2500 5000
microns^2 : pollen
0
C
HA
RC
O
A
L
CHARGE POND
Plymouth County, MA
FOSSIL POLLEN AND CHARCOAL
1981
analyst: A.E. Backman
CHARGE POND
Plymouth County, MA
FOSSIL POLLEN AND CHARCOAL
1981
14	 Fire 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
DEPTHINSEDIMENT(mm)
ca. 1760 AD
ca. 1700 AD
20
%
0
SP
RUC
E
20
%
0
PINE
20 40
%
0
BIRC
H
%
0
O
A
K
20 40
%
0
ALD
ER
%
0
G
R
ASSES
20
%
0
AG
RICU
LTU
RA
L
IN
DICATO
R
S
200 400 600
microns^2 : upland pollen
0
C
H
AR
CO
AL
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
15Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
DEPTHINSEDIMENT(cm)
6240 ± 110
5960 ± 120
4930 ± 90
3830 ± 100
3090 ± 90
2370 ± 100
1970 ± 80
ca. 1760 AD
C-14AGE(BP)
20
%
0
SPR
UCE
20 40
%
0
HEM
LO
CK
20 40
%
0
W
H
ITE
PIN
E
20 40
%
0
BIRC
H
20
%
0
O
AK
20
%
0
ALD
ER
%
0
G
RASSES
%
0
AG
RICU
LTURA
L
IND
IC
ATO
RS
200 400 600
microns^2 : pollen
0
CH
ARCO
AL
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
16	 Fire 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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20	 Fire 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; 2
USDA 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,
21Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
Site name State Forest
Type
Trees and forest type Pre-Euro-fire
interval
Post- Eurofire
interval
Site 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
23Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
24	 Fire 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.
25Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
26	 Fire 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.
27Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
28	 Fire 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
Topo-
resistance
Roughness
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
29Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
31Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
32	 Fire 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 0.4o
C 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).
33Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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®
ArcGIS™
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 95percent
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
41Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
42	 Fire 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
43Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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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49Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 ft2
ac-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; 2
USDA Forest Service, Northern Research
Station, 1992 Folwell Ave., St. Paul, MN 55108; 3
USDA
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.
51Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
Table1.—AreaoftimberlandintheEasternUnitedStates,inmillionsofacresandfromthemostrecent
inventory,byregionandbroadforesttype,withpercentofregiontotal1
1
Source:ForestInventoryDatabase(Miles2005).
a
Includesoak-hickoryandoak-pineforest-typegroups.
b
Includesswampchestnutoak-cherrybarkoak,sweetgum-Nuttalloak-willowoak,andovercupoak-waterhickoryforesttypes.
c
Michigan,Minnesota,andWisconsin.
d
Illinois,Indiana,Iowa,Missouri.
e
Connecticut,Massachusetts,Maine,NewHampshire,RhodeIsland,andVermont.
f
Delaware,Maryland,NewJersey,NewYork,Ohio,Pennsylvania,andWestVirginia.
g
Florida,Georgia,Kentucky,NorthCarolina,SouthCarolina,andVirginia.
h
Alabama,Arkansas,Louisiana,Mississippi,Oklahoma,Tennessee,andTexas.
i
Kansas,Nebraska,NorthDakota,andSouthDakota.
TotalUplandoaka
Otherupland
deciduousLowlandoakb
Otherlowland
deciduousConiferOther
AreaPercentAreaPercentAreaPercentAreaPercentAreaPercentAreaPercent
LakeStatesc
49.088.301724.9051-03.978%11.39230.521
CentralStatesd
25.2119.06762.0280.0902.96120.8530.231
NewEnglandStatese
31.314.521416.5853-00.8139.30300.100
Mid-AtlanticStatesf
56.0326.934822.50400.1102.2143.8270.461
Atlanticstatesg
96.0147.84501.0413.9149.601032.99340.621
Gulfstatesh
105.8552.05490.1008.03810.131035.00330.531
Plainsstatesi
5.331.56290.55100.0101.32251.67310.234
Total368.82160.274367.71812.15330.98895.03262.691
52	 Fire 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.
53Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
Table2.—Timberlandareaofuplandoakforesttypegroupsbyforesttypeandstandsizeclass,forthestateseastoftheGreatPlains
ForesttypeTotalSeedling-saplingPoletimberSawtimberNonstocked
Area
Percentof
TotalArea
Percentof
TotalArea
Percentof
TotalArea
Percentof
Total
Oak/pinegroup6.90.7104.3621.928--
Whitepine/redoak/whiteash3,836.90420.6111,175.70312,240.6058--
Easternredcedar/hardwood2,895.70769.2271,481.5051644.922--
Longleafpine/oak1,248.90639.651330.726275.9222.70
Shortleafpine/oak4,398.20579.9131,438.80332,379.5054--
Virginiapine/southernredoak2,178.90439.120820.538919.342--
Loblollypine/hardwood15,792.106,104.70393,722.50245,959.80385.10
Slashpine/hardwood1,797.3069739441.725653.53650
Otherpine/hardwood2,846.60707.225978.7341,157.80412.90
Totaloak/pinegroup35,001.5010,358.003010,394.403014,233.204015.700
Oak/hickorygroup1,444.10922.564243.717277.919--
Postoak/blackjackoak5,680.30808.5142,460.90432,410.9042--
Chestnutoak5,728.70165.431,750.10313,813.2067--
Whiteoak/redoak/hickory49,419.705,586.101113,927.102829,906.4061--
Whiteoak4,779.1072.621,183.10253,523.5074--
Northernredoak3,782.20110.53805.5212,866.2076--
Yellow-poplar/whiteoak/redoak7,450.90739.7101,674.50225,036.6068--
Sassafras/persimmon965299.33139040275.729--
Sweetgum/yellow-poplar6,649.202,614.80391,798.60272,235.8034--
Buroak582.126.75118.220437.275--
Scarletoak527.630.96207.939288.855--
Yellow-poplar1,250.20176.51437830695.656--
Blackwalnut653.881.512186.529385.959--
Blacklocust474.9168.235206.94499.821--
Southernscruboak1,281.10904.871326.62539.5310.21
Chestnutoak/blackoak/scarletoak3,051.701776944.4311,930.4063--
Redmaple/oak2,381.6036815965.1411,048.5044--
Mixeduplandhardwoods29,166.607,640.70268,851.603012,652.104322.20
Totaloak/hickorygroup125,268.8020,893.701736,418.702967,924.005432.400
Total(Thousandacres)160,270.3031,251.701946,813.102982,157.205148.10
Total(Thousandhectares)64,861.4012,647.6018,945.3033,249.0019.5
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,
55Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
57Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
59Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
61Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
62	 Fire 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 theWisconsin 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
importanceoftheoaksforwildlifefoodandcover.Finally,sevenreasonsforsustainingoakforestsarepresented.
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
63Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
65Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
Oak-
pine
Oak-
hickory
Oak-gum-
Cypress
Oak types
totalYear
-------------------------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.
67Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 ¼ 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
69Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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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Davis, M.B. 1981. Quaternary history and the stability
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73Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
75Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
77Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
79Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
81Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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).
82	 Fire 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
83Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
85Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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; 2
USDA 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
91Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
92	 Fire 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
93Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
95Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
FuelLoading(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 100hour
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
FuelLoading(Mg/ha)
0
10
20
30
40
50
60
70
80
Xeric - 6% Decomposition
Mesic - 8% Decomposition
97Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
99Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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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104	 Fire 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.
105Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
106	 Fire 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
107Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 CaCO3
825-898b
Mg MgCO3
900b
Mg base metal 1107b
P potassium
hexametaphosphate
1320b
Ca base metal 1487b
a
Agee (1993).
b
Weast (1969).
108	 Fire 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.
109Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
NO3
and
NH4
+
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
110	 Fire 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 NO3
and
PO4
3also
were greater in
leachate from burned sites, and the degree of enrichment
111Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
PO4
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
113Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
114	 Fire 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.
115Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
116	 Fire 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 PO4
3B
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
117Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 ↓
118	 Fire 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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123Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
totherelativecompetitivenessofoakreproductionbenefitingfromorbeingharmedbyprescribedfires.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; 2
USDA Forest Service, Northeastern Research
Station, Parsons, WV 26287; 3
Connecticut Agricultural
Experiment Station, Department of Forestry and Horticulture,
New Haven, CT 06504; †
Corresponding author, 814-563-1040,
email: pbrose@fs.fed.us.
124	 Fire 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
ReviewedStudies(%)
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.
125Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
126	 Fire 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
ReviewedStudies(%)
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.
127Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
128	 Fire 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
129Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
130	 Fire 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.
Literature Cited
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forests. Bioscience. 42: 346-353.
Adams, A.S.; Rieske, L.K. 2001. Herbivory and fire
influence white oak. Quercus alba L. seedling
vigor. Forest Science. 47(3): 331-337.
Apsley, D.K.; McCarthy, B.C. 2004. White-tailed deer
herbivory on forest regeneration following fire
and thinning treatments in southern Ohio mixed
oak forests. In Yaussy, D.A.; Hix, D.M.; Long,
R.P.; Goebel, P.C, eds. Proceedings, 14th central
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hardwood forest conference. Gen. Tech. Rep. NE-
316. Newtown Square, PA: U.S. Department of
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136	 Fire 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.
137Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 (64o
F) (Iverson and
Hutchinson 2002). Flinn and Pringle (1983) found
that rhizome death of several forest understory species
occurred only as temperatures approached 60o
C (140o
F).
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
138	 Fire 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.
139Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
140	 Fire 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.
141Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
142	 Fire 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.
143Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
144	 Fire 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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148	 Fire 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
149Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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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151Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
Fire - Fauna Interactions
152	 Fire 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.
153Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
154	 Fire 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
155Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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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158	 Fire 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.
159Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
160	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
Table1.—DetailsoffiresandamphibianandreptileresponsestothosefiresintheeasternoakforestsoftheUS.
Locationand
reference
Typeoffire
treatments
FireintensityFuelconsumptionPatchinessof
consumption
SpeciesevaluatedAnimalr80esponse
Indiana(Palis
1995)
Aprilburn;Prescribed
burnsetat1615hrs
NighttimefireNAa
80percentof
1,000acres
burned
Amphibiansand
reptilesofthatregion
Nodirectmortality
observed;resulting
amphibianand
reptilenumbersand
assemblageappeared
typical
Pennsylvania
(Kirklandand
others1996)
BurnedinNovemberCharredtree
trunks;killed
manysaplings;
charredvegetation
ingroundand
shrublayers
NANAAmericantoadMoretoadsonburned
thanunburnedsites
NorthCarolina
(Fordandothers
1999)
BurnedinAprilHighintensity
prescribedfire
NANAWoodland
salamanders
Nodifferencein
salamanderabundance
detectedbetween
burnedandunburned
sites
SouthCarolina
Piedmont(Floyd
andothers2002)
Firstroundoffires
inFebruary-March
1999;striphead
firesset15to30feet
apart;secondround
offiresinApril2000;
stripheadfiresset10
to20feetapart;
Firstroundaverage
flamelengthwas
>1foot;second
roundaverage
flamelength<1
foot
NANATotalherpetofauna,
frogs,toads,
salamanders,newts,
lizards,turtles,snakes
Nodifferencein
abundance,diversity,
andevenessintotal
herpetofaunaorby
taxonomicgroups
161Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
Georgiaupper
CoastalPlain
(Moseleyand
others2003)
Winterburns;stands
hadbeenburned
every2-3yearsinthe
previous9years
LowintensityMorelitterin
unburned;more
bareground
inburned;no
differenceincoarse
woodydebris
volumebetween
burnedand
unburned
NATotalherpetofauna;
amphibians,anurans,
salamanders,reptiles
Nodifferencein
overallamphibianand
reptile,salamander,
oranuranabundance,
richness,anddiversity
betweenburnedand
unburned;reptile
abundanceand
diversityweregreater
inburnedthan
unburnedsites
Virginia
Piedmont
(Keyserand
others2004)
Seasonalprescribed
burns(winter,
spring,summer)
appliedtofirst-stage
shelterwood-harvested
stands(to48ft2
basal
areaperacre)cut3to
5yearspriortoburns
Averagefire
temperature39
inchesabovethe
groundwas526
°Finwinterburn,
648°Finspring
burn,and485°F
insummerburn
Allfiresreduced
fine,medium,
coarsefuelloads
(finefuelswere
eliminated;coarse
fuelswerepartially
reduced),yetduff
layerremained
intact
Springand
summerburns
causedashift
fromshrub
dominatedto
herbaceous
dominated
understory
Allamphibiansand
reptiles,especially
easternred-backed
salamanders,
Americantoads,
northernfence
lizards(Sceloporus
undulatus),little
brownskinks,and
southeasternfive-
linedskinks(Eumeces
inexpectatus)
Nodifference
inabundanceof
salamanders,toads,
andallamphibians
betweencontrols
anddifferent
seasonalburns;lizard
abundancewas
greaterinburnedthan
unburnedstands
aNotAvailable
Table1.—continued.
162	 Fire 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.
163Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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).
164	 Fire 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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167Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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; 2
USDA
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.
168	 Fire 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
169Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
170	 Fire 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
171Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
172	 Fire 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)
173Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
174	 Fire 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
175Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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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180	 Fire 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; 2
USDA Forest Service, Northern Research
Station, Box 404, Parsons, WV 26287; †
Corresponding author,
304-392-1606, email: pdk2@meadwestvaco.com.
181Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 nonvolant
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
182	 Fire 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
183Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
184	 Fire 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.
185Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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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191Fire 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
192	 Fire 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.
193Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
194	 Fire 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
195Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
196	 Fire 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
197Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
literature cited
Blanchard, B.; Ryan, R.L. 2004. Community
perceptions of wildland fire risk and fire hazard
reduction strategies at the wildland-urban interface
in the Northeastern United States. In: Murdy,
James, comp., ed. Proceedings of the 2003 northeast
recreation research symposium; Bolton Landing,
NY. Gen. Tech. Rep. NE-317. Newtown Square,
PA: U.S. Department of Agriculture, Forest Service,
Northeastern Forest Experiment Station: 285-294.
Blanchard, B.P. 2003. Community perceptions of
wildland fire risk and fire hazard reduction
strategies at the wildland-urban interface in the
Northeastern United States. Thesis. University of
Massachusetts, Amherst.
Bright, A.; Carroll, J. 2004. An Assessment of public
perceptions of fuel reduction activities on National
Forests. Fort Collins, CO: Department of Natural
Resource Recreation & Tourism, Report to North
Central Research Station. U.S. Department of
Agriculture, Forest Service. 92 p.
Brunson, M.W.; Evans, J. 2005. Badly burned?
Effects of an escaped prescribed burn on social
acceptability of wildland fuels treatments. Journal
of Forestry. April/May: 134-138.
Carpenter, E.H; Taylor, J.G; Cortner; H.J.; Gardner;
P.D.; Zwolinski, M.J.; Daniel, T.C. 1986. Targeting
audience and content for forest fire information
programs. The Journal of Environmental Education.
17(3): 33-41.
Cortner, H.J.; Zwolinski; M.J.; Carpenter, E.H.; Taylor,
J.G. 1984. Public support for fire-management
policies. Journal of Forestry. 82(6): 359-360.
Gardner, P D.; Cortner, H.J. 1988. An assessment
of homeowner’s perceptions of wildland fire
hazards: A case study from southern California.
In: Whitehead, Emily B., ed. Arid Lands Today and
Tomorrow, Boulder: Westview Press.
Loomis J.B.; Bair, L.S.; Gonzalez-Caban, A. 2001.
Prescribed fire and public support: knowledge
gained, attitudes changed in Florida. Journal of
Forestry. 99: 11, 18-22.
McCaffrey, S.M. 2005. Understanding public
conceptions of wildfire risk. In: Martin, W.; Raish,
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C.; Kent, B., eds. Wildfire risk: human perceptions
and management implications, Resources for The
Future Press.
McCaffrey, S.M. 2002. For want of defensible space a
forest is lost: Homeowners and the wildfire hazard
and mitigation in the residential wildland intermix
at Incline Village, Nevada. Dissertation, University
of California, Berkeley.
Monroe, M.C.; Pennisi, L.; McCaffrey, S.M.; Mileti, D.
2005. Social science to improve fuels management:
a synthesis of research related to communicating
with the public on fuels management efforts. Gen.
Tech. Rep. NC-267. St Paul, MN: U.S. Department
of Agriculture, Forest Service, North Central Research
Station. 42 p.
Shelby, Bo; Speaker, Robert W. 1990. Public attitudes
and perceptions about prescribed burning. In:
Walstad, John D.; Radosevich, Steven R.; Sandberg,
David V., eds. Natural and prescribed fire in Pacific
northwest forests. Corvallis, OR: Oregon State
University Press: 253-259.
Shindler, Bruce; Leahy, J.; Toman, E. 2003. Public
acceptance of forest conditions and fuel reduction
practices: a survey of citizens in communities
adjacent to national forests in Minnesota,
Wisconsin, and Michigan. Report to North Central
Research Station, USDA Forest Service and the Joint
Fire Science Program.
Shindler, B.; Toman, E. 2003. Fuel reduction strategies
in forest communities: A longitudinal analysis.
Journal of Forestry. 101(7): 9-15.
Shindler, Bruce; Reed, Michelle; Kemp, Beth; McIver,
James. 1996. Forest management in the Blue
Mountains: Public perspectives on prescribed fire
and mechanical thinning. Corvallis, OR: Dept of
Forest Resources, Oregon State University.
Slovic, P.B. Fischhoff; Lichtenstein, S. 1987. Behavioral
decision theory perspectives on protective
behavior. In: Weinstein, Neil, ed. Taking care:
understanding and encouraging self-protective
behavior. New York: Cambridge University Press:
14-41.
Weisshaupt, B.R.; Carroll, M.S.; Blatner, K.A.;
Robinson, W.D.; Jakes, P.J. 2005. Acceptability
of smoke from prescribed forest burning in the
northern inland west: a focus group approach.
Journal of Forestry. 103(4): 189-193.
Winter, Greg; Vogt, C.; McCaffrey, S.M. 2005.
Community views of fuels management on the
Mark Twain National Forest and comparisons
to other study sites. Survey Data Report. Prepared
for North Central Research Station, USDA Forest
Service.
Winter, Greg; Vogt, C.; McCaffrey, S.M. 2004.
Examining social trust in fuels management
strategies. Journal of Forestry. 102(6): 8-15.
Winter, G.; Vogt, C.; Fried, J.S. 2002. Demographic
and geographic approaches to predicting public
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Station, USDA Forest Service.
199Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 theycan 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; 2
USDA Forest Service, Region 9, Fire and
Aviation Management, 27750 State Route 7, Marietta, OH
45750; 3
USDA Forest Service, Monongahela National Forest,
200 Sycamore St., Elkins, WV 26241; †
Corresponding author,
517-355-7740 ext. 105, email: jcharney@fs.fed.us.
200	 Fire 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.
201Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
Figure 2.—Population density in the eastern United States.
Figure 3.—Locations of Interstate Highways overlaying population density in the eastern United
States.
202	 Fire 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.
203Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
204	 Fire 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).
205Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
206	 Fire 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
207Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
208	 Fire 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.
REFERENCES
Achtemeier, G.L. 2001. Simulating Nocturnal Smoke
Movement. Fire Management Today. 61: 28-33.
Breyfogle, S.; Ferguson, S.A. 1996. User assessment of
smoke-dispersion models for wildland biomass
burning. Gen. Tech. Rep. PNW-379. Portland,
OR: U.S. Department of Agriculture, Forest Service,
Pacific Northwest Research Station. 30 p.
Hardy, C.; Ottmar, R.D.; Peterson, J.; Core, J. 2002.
Smoke management guide for prescribed and
wildland fire—2000 edition. PMS 420-2. NFES
Figure 11.—BlueSky output using the weather conditions on April
24th, 2005 and a hypothetical distribution of prescribed burns
across the Great Lakes, Ohio Valley, and southern Canada.
209Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
1279. Boise, ID: National Wildfire Coordination
Group. 226 p.
Hicks, R.R. 1997. A resource at the cross-roads: a
history of the central hardwoods. In: Pallardy,
Stephen G.; Cecich, Robert A.; Garrett, H. Eugene;
Johnson, Paul S., eds. Proceedings of the 11th central
hardwood forest conference; 1997 March 23-26;
Columbia, MO. Gen. Tech. Rep. NC-188. St. Paul,
MN: U.S. Department of Agriculture, Forest Service,
North Central Forest Experiment Station: 1-2.
Lavdas, L.G. 1996. Program VSMOKE-Users manual.
Gen. Tech. Rep. SRS- 6, Asheville, NC: U.S.
Department of Agriculture, Forest Service, Southern
Research Station. 147 p.
Midwest RPO. 2001. Regional haze and visibility in
the upper Midwest. Midwest Regional Planning
Organization, Technical Report, (http://www.ladco.
org/tech/monitoring/docs_gifs/rhreport.pdf ).
O’Neill, S.M.; Ferguson S.A.; Peterson J.; Wilson, R.
2003. The BlueSky Smoke Modeling Framework
(www.blueskyrains.org), In: Proceedings, fifth
symposium on fire and forest meteorology. Orlando,
FL: American Meteorological Society.
Sestak, M.L.; Riebau, A.R. 1988. SASEM, Simple
Approach Smoke Estimation Model. Tech. Note
382. U.S. Bureau of Land Management. 31 p.
Stringer, J.W.; Loftis, D.L. 1999. Forward to the 12th
central hardwood forest conference. In: Stringer,
Jeffrey W.; Loftis, David L.; Lacki, Michael; Barnes,
Thomas; Muller, Robert A., eds. Proceedings of
the 12th central hardwood forest conference; 1999
March 1-2; Lexington, KY. Gen. Tech. Rep. SRS-
024. Asheville, NC: U.S. Department of Agriculture,
Forest Service, Southern Research Station: ii.
U.S. Bureau of the Census 2000. 2000 U.S. Census
Data. November, 2005: http://venus.census.gov/
cdrom/lookup.
U.S. EPA 2003. Guidance for Estimating Natural
Visibility Conditions Under the Regional Haze
Program. EPA-454/B-03-005.
U.S. EPA 2006. Nonattainment Area Map. Green
Book. Updated 09/2005. Available at: http://www.
epa.gov/oar/oaqps/greenbk/mapnpoll.html
U.S. EPA 1995. User’s Guide for the Industrial Source
Complex (ISC3) Dispersion Models. Office of Air
Quality Planning and Standards, Research Triangle
Park, NC, EPA-454/B-95-003a.
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
D.A.; Hix, D.M.; Long, R.P.; Goebel, P.C., eds.
Proceedings, 14th central hardwood forest conference;
Wooster, OH. Gen. Tech. Rep. NE-316. Newtown
Square, PA: U.S. Department of Agriculture, Forest
Service, Northeastern Research Station:193-199.
Wickman, T.; Acheson, A. 2005. Fuels planning:
science synthesis and integration; environmental
consequences fact sheet 11: Smoke Impact
Spreadsheet (SIS) Model. Res. Note. RMRS-RN-
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Agriculture, Forest Service, Rocky Mountain Research
Station. 2 p.
210	 Fire 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; 2
Rocky Mountain Research
Station, Forestry Sciences Laboratory, 800 East Beckwith,
Missoula, MT 59801; †
Corresponding author, 603-868-7624,
email: ktsmith@fs.fed.us.
211Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
213Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
214	 Fire 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.
215Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
216	 Fire 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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218	 Fire 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.
219Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
220	 Fire 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.
221Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
223Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
224	 Fire 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
225Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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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University Press: 269-279.
232	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
White, R.H.; Weise, D.R.; Mackes, K.; Dibble, A.C.
2002. Cone calorimeter testing of vegetation: An
update. In: Proceedings of the 35th
international
conference on fire safety, 17th
international conference
on thermal insulation, and 9th
international
conference on electrical and electrical products;
2002 July 22-24; Columbus, OH. Sissonville, WV:
Products Safety Corp.
Winberry, J.J.; Jones, D.M. 1973. Rise and decline
of the “miracle vine:” kudzu in the southern
landscape. Southeastern Geographer. 13: 61-70.
Wolters, G.L.; Sieg, C.H.; Bjugstad, A.J.; Gartner, F.R.
1994. Herbicide and fire effects on leafy spurge
density and seed germination. Res. Note RM-526.
Fort Collins, CO: US Department of Agriculture,
Forest Service, Rocky Mountain Forest and Range
Experiment Station. 5 p.
233Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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; 3
Heritage Preserves, South Carolina Department
of Natural Resources,1000 Assembly Street, Columbia, SC
29201; 4
Florida 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.
234	 Fire 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).
235Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
236	 Fire 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
237Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
238	 Fire 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;
239Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
240	 Fire 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
241Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
242	 Fire 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
243Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
244	 Fire 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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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
250	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
251Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
252	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
(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.
253Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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;
254	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
255Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
256	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
257Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
258	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
259Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
260	 Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
	 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
261Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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
262	 Fire 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.
263Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
264	 Fire 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.
265Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
266	 Fire 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.
267Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
Poster Abstracts
268	 Fire 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
269Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
THERMOCOUPLE PROBES IN SURFACE FIRES
Anthony S. Bova1†
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 2
Ecologist, Northern Research Station;
†
Corresponding author, 359 Main Road, Delaware, OH 43015,
740-368-0093, email: abova@fs.fed.us.
270	 Fire 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; 2
Professor, 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.
271Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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, 2
Northern Research
Station, USDA Forest Service, East Lansing, MI 48823 †
Corresponding
author; 609-894-0325, email: kennethclark@fs.fed.us.
272	 Fire 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 NO3
-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; 2
Professor, Department of Forestry and Natural
Resources, Clemson University; 3
Research Forester, USDA Forest Service,
Southern Research Station; 4
Professor, Department of Entomology, Soils,
and Plant Sciences, Clemson University; 5
Department Chair, Department
of Applied Economics and Statistics, Clemson University; †
Corresponding
author, 265 Lehotsky Hall, Clemson, SC 29634, 864-656-3054, email:
coatest@clemson.edu.
273Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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; 2
Fire Ecologist, USDA Forest
Service, Southern Region; 3
Biological Science Technician (Fire Effects),
North Cascades National Park; 4
Fire Management Officer (ret.), Great
Smoky Mountains National Park; 5
Fire Management Officer, Great
Smoky Mountains National Park; †
Corresponding author, P.O. Box 645,
Panguitch, UT 84759, (435) 676-9372, email: dcohen@fs.fed.us.
274	 Fire 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. Bova2
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 2
Physical Scientist, USDA Forest Service, Northern Research
Station; †
Corresponding author, 359 Main Road, Delaware, OH 43015,
740-368-0093, email: mbdickinson@fs.fed.us.
275Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
276	 Fire 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.
277Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
278	 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
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; 2
Professor 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.
279Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
280	 Fire 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; 2
Assistant 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.
281Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
ESTIMATING FIRE BEHAVIOR FROM OVERHEAD IMAGERY IN EASTERN
OAK FORESTS
Robert Kremens1†
, Anthony Bova2
, 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; 2
Physical
Scientist, USDA Forest Service, Northern Research Station; 3
Ecologist,
USDA Forest Service, Northern Research Station; 4
Integration Engineer,
Rochester Institute of Technology; 5
Graduate Student, Rochester Institute
of Technology; †
Corresponding author, 54 Lomb Memorial Drive,
Rochester, NY 14623, 585-475-7286, email: kremens@cis.rit.edu.
282	 Fire 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; 2
Professor 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.
283Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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-m2
plots 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; 2
Ohio State University
Extension; †
Corresponding author, 359 Main Road, Delaware, OH
43015-8640, 740-368-0050, email: rlong@fs.fed.us.
284	 Fire 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; 2
Former
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.
285Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 ™
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.
286	 Fire 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, 2
Graduate 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.
287Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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; 2
Assistant 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.
288	 Fire 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.
289Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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; 2
Research Ecologist, USDA Forest Service,
Northeastern Research Station; 3
Dendrochronologist, USDA Forest
Service, Northeastern Research Station; 4
Professor 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.
290	 Fire 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; 2
Geographer, USDA Forest
Service Pacific Southwest Research Station; 3
Professor, 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.
291Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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.
292	 Fire 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; 2
University System Forest Manager, Arkansas Forest
Resources Center; 3
George H. Clippert Endowed Chair, University of
Arkansas – Monticello, School of Forest Resources; 4
Forestry Instructor,
University of Arkansas, Cooperative Extension Service; †Corresponding
author, P.O. Box 3468, Monticello, AR 71656, 870-460-1052, email:
montgomeryr@uamont.edu.
293Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 2
Regional 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.
294	 Fire 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; 2
Assistant 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.
295Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
USE OF TEMPERATURE-SENSITIVE PAINTS AS AN INDEX OF HEAT
OUTPUT FROM FIRE
Joanne Rebbeck1†
, Daniel Yaussy2
, Louis Iverson3
, Todd Hutchinson3
, Robert Long4
, Anthony Bova5
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, 2
Forester, 3
Ecologist,4
Plant Pathologist, 5
Physical
Scientist, and 6
Ecologist, USDA Forest Service, Northern Research
Station; †
Corresponding author, 359 Main Road, Delaware, OH, 43015,
740-368-0054, email: jrebbeck@fs.fed.us
296	 Fire 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.
297Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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; 2
Professor 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.
298	 Fire 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; 2
Wildlife 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.
299Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 firevegetation
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.
301Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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 360o
F (range: 142 to 696o
F). 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 > 300o
F compared with only 5 percent for stems exposed to temperatures < 300o
F.
1
Chief Scientist, The Connecticut Agricultural Experiment Station,
Department of Forestry and Horticulture; 2
Forester, 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.
302	 Fire 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; 2
Professor and Extension Specialist, Rangeland Ecology
and Management, OSU; 3
Biologist, US Fish & Wildlife Service;
†
Corresponding author, 4922 South Coyle Road, Stillwater, OK 74074,
405-743-4714, email weir@okstate.edu.
303Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference	 GTR-NRS-P-1
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, 2
Ecologists, and 3
Plant 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.
www.nrs.fs.fed.us
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