United States Department of Agriculture

Forest 

Service

General Technical 

Report NRS-135

Northern 

Research Station July 2014

The Fire–Oak Literature of 

Eastern North America: 

Synthesis and Guidelines

Patrick H. Brose

Daniel C. Dey

Thomas A. Waldrop
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Abstract

Guidelines for using prescribed fire to regenerate and restore upland oak forests, 

woodlands, and savannas in eastern North America were developed by synthesizing the 

results of more than 100 scientific publications. The first four chapters provide background 

information on the values of oak ecosystems, eastern fire history, oak’s adaptations to fire, 

and the findings of fire-oak research conducted over the past 50 years. The final chapter 

synthesizes that background information into guidelines that explain how to use prescribed 

fire to facilitate oak seedling establishment, release oak reproduction from competing 

mesophytic hardwoods, and rehabilitate open oak woodlands, oak savannas, and scrub oak 

communities. A reference section is also provided for readers desiring to delve more deeply 

into the associations between periodic fire and oak forests, woodlands, and savannas.

The Authors

PATRICK H. BROSE is a research forester with the U.S. Forest Service, Northern 

Research Station, Irvine, PA.

DANIEL C. DEY is a research forester and project leader with the U.S. Forest Service, 

Northern Research Station, Columbia, MO.

THOMAS A. WALDROP is a research forester and team leader with the U.S. Forest 

Service, Southern Research Station, Clemson, SC.

Cover Photos

Photos illustrating two of the roles that periodic prescribed fire can fulfill in the sustaining of 

mixed-oak forests in eastern North America. Upper left: oak forest succeeding to a mixed 

mesophytic forest in the absence of periodic fire. Upper right: seedbed preparation burn in 

a mature oak stand. Lower left: release burn in an oak shelterwood. Lower right: oak stand 

with abundant, vigorous oak reproduction resulting from the use of prescribed fire. Photos 

by Patrick Brose and Daniel Yaussy, U.S. Forest Service.

Manuscript received for publication 27 December 2013

This synthesis was funded by the Joint Fire Science Program
The Fire—Oak Literature of Eastern North America: 

Synthesis and Guidelines

Patrick H. Brose

Daniel C. Dey

Thomas A. Waldrop

General Technical Report NRS-135

Quality Assurance

This publication conforms to the Northern Research Station’s Quality Assurance Implementation Plan 

which requires technical and policy review for all scientific publications produced or funded by the Station. 

The process included a blind technical review by at least two reviewers, who were selected by the 

Assistant Director for Research and unknown to the author. This review policy promotes the Forest Service 

guiding principles of using the best scientific knowledge, striving for quality and excellence, maintaining 

high ethical and professional standards, and being responsible and accountable for what we do.
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Contents

CHAPTER 1: INTRODUCTION.....................................................................................1

The Values of Oak Ecosystems ...............................................................................2

The Fire-Oak Synthesis Project ...............................................................................7

CHAPTER 2: FIRE HISTORY OF EASTERN NORTH AMERICA .................................8

Paleoecology Studies ..............................................................................................8

Dendrochronology Studies .................................................................................... 10

Synthesis ...............................................................................................................14

CHAPTER 3: ADAPTATIONS OF OAK TO FIRE.........................................................17

Acorns ...................................................................................................................17

Seedlings...............................................................................................................18

Saplings.................................................................................................................21

Mature Trees ..........................................................................................................22

Synthesis ...............................................................................................................24

CHAPTER 4: REVIEW OF FIRE EFFECTS STUDIES ...............................................26

Mature Stands .......................................................................................................27

Young Stands.........................................................................................................34

Immature Stands ...................................................................................................35

Synthesis ...............................................................................................................36

CHAPTER 5: GUIDELINES FOR USING FIRE IN OAK ECOSYSTEMS....................41

Essential Background Information .........................................................................41

Seedbed Preparation Burning ...............................................................................46

Release Burning ....................................................................................................51

Ecological Restoration Burning..............................................................................63

EPILOGUE ..................................................................................................................73

ACKNOWLEDGMENTS ..............................................................................................73

LITERATURE CITED................................................................................................... 74

APPENDIX 1: COMMON AND SCIENTIFIC NAMES OF THE FLORA AND 

FAUNA MENTIONED IN THIS REPORT.....................................................................98
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1

CHAPTER 1: INTRODUCTION

Of all the plant communities that make up the eastern hardwood biome, the oak1 ecosystems 

(forests, savannas, shrublands, and woodlands) are perhaps the most important because of their 

vastness, diversity, and many ecological and economic values (Smith 2006). At least 50 oak 

species grow east of the 100th meridian, and some type of oak community is found in every 

state and adjacent Canadian province (Burns and Honkala 1990, Stein et al. 2003). Some oak 

species such as white oak are widespread and occur in most states and provinces, while others 

like Mohr oak are restricted to a few counties in Oklahoma and Texas. Oaks occur as solitary 

individuals in grassland environments, as a few scattered trees in northern hardwood forests, as 

substantial portions of complex, mixed-species assemblages, and as nearly pure monocultures 

(Fig. 1). They vary in size from the shrubby runner oak and vine-like dwarf live oak to the 

massive bur oak and cherrybark oak. Oaks grow in a wide array of habitats, from seasonallyflooded bottomlands, to rich mountain coves, to xeric ridges and sandhills. Their widespread 

occurrence, variation in size and form, and diversity of habitats are three of the reasons the 

generic oak was designated by Congress as America’s national tree (Arbor Day Foundation 

2001, 2004).

1 Scientific names of flora and fauna are listed in the Appendix.

Figure 1.—Oaks occur as solitary trees (upper left), scattered individuals in other forest types (upper right), substantial portions 

of mixed-species stands (lower left), and as nearly pure monocultures (lower right). Photos by Patrick Brose, U.S. Forest Service.
2

Despite this widespread occurrence and dominance, oak ecosystems are declining in abundance 

and require careful management to be sustained. The scientific literature abounds with studies 

reporting chronic oak regeneration failures and the replacement of oaks by mesophytic 

hardwood species (Abrams and Downs 1990, Aldrich et al. 2005, Healy et al. 1997, Schuler 

and Gillespie 2000, Woodall et al. 2008). Pennsylvania is a prime example of the oak 

regeneration problem. Recent U.S. Forest Service Forest Inventory and Analysis data show 

that for all stems larger than 2 inches in diameter at breast height (d.b.h.), red maple and 

sweet birch are the two most common species, and the most common oak, northern red, is the 

ninth most abundant hardwood (McWilliams et al. 2004). The disparity is especially evident 

in the sapling (1- to 5-inch d.b.h.) and pole (6- to 10-inch d.b.h.) size classes where red maple 

outnumbers all oaks by ratios of 6:1 and 3:1, respectively (Fig. 2). As mature oaks succumb to 

various causes, they are being replaced by hardwood species other than oaks. This conversion is 

highly undesirable for many ecological and economic reasons.

The Values of Oak Ecosystems

One of the primary reasons for retaining and promoting oak is its importance to wildlife, and 

that value begins with the foliage. Oaks produce copious amounts of foliage, and these leaves 

are the base of the food chain in the eastern hardwood biome. Among the genera of woody 

plant species native to the mid-Atlantic region, oak ranks first in the ability to support native 

butterfly and moth (Lepidoptera) species (Fig. 3) (Tallamy and Shropshire 2009). The authors 

documented that the larva of 534 native lepidopteran species consume oak leaves. These larvae 

serve as food for other insects, birds, and small mammals (Fig. 4). Birch and maple, common 

oak-replacement trees, support considerably fewer native lepidopteran species. In addition to 

foliage, oak bark supports insects of all types because the rough texture provides refuge from 

Figure 2.—In many parts of eastern North America, oak forests are being replaced by 

various mesophytic hardwood species such as red maple. Photo by Patrick Brose, U.S. 

Forest Service.
3

Figure 3.—The orangestriped oakworm is one of the 534 native lepidopteran 

species that feed on oak foliage. Photo by Kurt Gottschalk, U.S. Forest Service.

Figure 4.—Cerulean warblers (top left), scarlet 

tanagers (top right), and red-eyed vireos 

(bottom left) are three neotropical migrant 

songbirds that make extensive use of oak 

ecosystems and the caterpillars found there. 

Photos by Scott Stoleson, U.S. Forest Service.
4

predators and weather compared to the smooth bark of birches and maples which provide little, 

if any, protection (Fig. 5). When comparing bird densities between mature stands dominated 

by oak or red maple, Rodewald and Abrams (2002) found that oaks support a significantly 

higher abundance of birds in all seasons. In addition, avian species richness is significantly 

greater in oak than in maple stands. Differences are greatest in autumn during masting and 

in spring when oak flowers attract insects. They predict that a conversion of oak forests to red 

maple would have a severe impact on bird communities in the eastern United States. In aquatic 

environments, oak leaves are a valuable source of detritus that directly and indirectly benefits 

numerous organisms (Rubbo and Kiesecker 2004).

The longevity and size of oaks are important to wildlife. Many of the oaks are long-lived, quite 

resistant to rot, and grow to large sizes. These characteristics indicate that, once established, 

oak forests are relatively stable plant communities (Smith 2006). Once oaks achieve canopy 

dominance, they likely will persist for decades to centuries as they readily withstand insect 

outbreaks, lightning strikes, wind events, and ice storms. When damage does occur, the ability 

to compartmentalize rot helps ensure their continued survival (Shigo 1984). When oaks are 

injured, the injury sites may become cavities which, in turn, become dens for wildlife. Dens are 

especially valuable as wildlife habitat when they occur in large trees (Godfrey et al. 2000).

Another structural characteristic that makes oaks important to wildlife is their broad, open 

crowns (Fig. 6). The canopies of oak forests and woodlands are usually uneven (vary in height 

and thickness), discontinuous (have small gaps between adjacent crowns), and contain large 

branches. This canopy heterogeneity is important habitat for bats and crown-nesting birds like 

Figure 5.—The rough bark of oak trees such as chestnut oak (left) serves as refuge for many insect species. 

The smooth bark of many mesophytic species such as red maple (right) lack this protective attribute. 

Photos by Patrick Brose, U.S. Forest Service.
5

the cerulean warbler. Hamel et al. (2004) reported that cerulean warblers nest and forage more 

in oak forests than mixed hardwood forests. Oak canopies are also easily penetrated by sunlight 

which allows the development of diverse herbaceous and shrub communities (Fig. 7) that 

provide further food and habitat for wildlife.

Figure 6. —The broad, open crowns of many mature oaks are important to bats and some 

neotropical bird species, especially in woodlands and savannas. Photo by Patrick Brose, 

U.S. Forest Service.

Figure 7. —The understory of many oak forests has the necessary light level to support a 

rich herbaceous plant community. Photo by Patrick Brose, U.S. Forest Service.
6

Acorns only occur intermittently but are a valuable food resource for many animals because 

they are relatively large, readily digestible, easily stored, and high in nutrition (Kirkpatrick 

and Pekins 2002, McShea and Healy 2002). Within the eastern hardwood biome, more than 

100 vertebrate species regularly consume acorns, ranging from important game animals such 

as white-tailed deer, black bear, tree squirrels, wild turkey, wood ducks, and ruffed grouse, 

to nongame species such as red-headed woodpeckers, blue jays, white-footed mice, and 

chipmunks. In the Appalachians and the Northeast, acorns are the primary source of hard mast 

since the loss of the American chestnut and the decline of American beech.

In oak ecosystems, many species of wildlife depend on acorns to such an extent that they exhibit 

physiological and population responses to fluctuations in acorn crops. For example, acorn 

availability can affect the weight, condition, reproductive rates, and antler characteristics of 

white-tailed deer (Wentworth et al. 1992). Similarly, rates of birth, survival, and dispersal in 

black bear vary with the abundance and distribution of acorn crops (Pelton 1989). Variation 

in mast production also influences population levels and distributions of a variety of small 

mammals and birds, including squirrels, jays, and woodpeckers (McShea 2000, Rodewald 

2003).

Oak species are of great economic importance to the eastern United States and Canada. The 

stumpage price for oak consistently ranks near or at the top among all commercial hardwood 

species. Oak saw logs are used for a variety of products ranging from cabinets, fine furniture, 

flooring, and whiskey barrels to dimension lumber, pallets, and railroad ties (Fig. 8). The woodproducts industry employs thousands of people in the eastern United States and Canada, a 

significant proportion of which is related to the growth, harvest, manufacturing, and delivery of 

products from oak.

Figure 8.—Oaks are renowned for their high 

quality timber (above) that serves as the raw 

material for a wide array of wood products (right). 

Photos by Patrick Brose, U.S. Forest Service.
7

The Fire-Oak Synthesis Project

The chronic, widespread oak regeneration problem is widely regarded by many natural resource 

professionals to be caused, at least in part, by the advent of fire exclusion policies in the early 

1900s (Abrams 1992, Brose et al. 2001, Nowacki and Abrams 2008). The pervasiveness of the 

oak regeneration problem and the role of fire exclusion in it have sparked tremendous research 

interest in the fire-oak association by scientists over the past 50 years. This research focuses on 

understanding the history of fire, especially before European settlement, and the effects of fire 

on hardwood species and other ecosystem attributes. All of this research has produced a wide 

array of scientific articles ranging from simple case studies in conference proceedings to complex 

studies published in peer-reviewed journals.

In 2010, the Joint Fire Science Program (JFSP) issued a request for proposals to synthesize the 

existing fire-oak literature. The ultimate goal was to develop guidelines for using prescribed 

fire to restore and sustain upland oak ecosystems in eastern North America (defined as east of 

the 100th meridian). This opportunity was especially interesting to us because we all have been 

involved in fire-oak research for much of our careers, publishing numerous scientific papers on 

the subject, and frequently giving oral presentations to a wide variety of audiences. This report 

is the culmination of that literature synthesis and one of the products of the JFSP project.

This report is organized around the fire-oak hypothesis (Abrams 1992, Arthur et al. 2012, Brose 

et al. 2001, Nowacki and Abrams 2008) which consists of four parts: 

1. Fire has been an integral part of upland oak ecosystems in eastern North America for 

millennia.

2. Oaks are superiorly adapted relative to other hardwoods to survive a periodic fire 

regime and exploit the postfire environment.

3. The cessation of a periodic fire regime in the early 1900s is a major cause for the 

current oak regeneration problem.

4. Prescribed fire can be used in some situations to help regenerate and restore upland oak 

ecosystems.

Chapter 2 addresses fire-oak hypothesis parts 1 and 3 by reviewing the fire history publications 

of the eastern hardwood biome. Chapter 3 covers part 2 of the fire-oak hypothesis and the 

silvical characteristics of oak that make it a superior survivor and competitor in a periodic fire 

regime. Chapter 4 reviews the published fire effects on oak reproduction and relates them to 

part 4 of the fire-oak hypothesis. Chapter 5 synthesizes the findings of these three chapters 

into guidelines on how to use prescribed fire in upland oak ecosystems to meet some common 

management objectives. An extensive reference section listing the papers used in this project is 

also provided.
8

CHAPTER 2: FIRE HISTORY OF EASTERN NORTH AMERICA

Parts 1 and 3 of the fire-oak hypothesis posit that periodic fire has been an integral disturbance 

in the upland oak ecosystems of eastern North America for millennia, and the sudden cessation 

of that fire regime is one of the major causes of the current oak regeneration problem (Abrams 

1992, Arthur et al. 2012, Brose et al. 2001, Nowacki and Abrams 2008). Both of these 

premises have been the focus of historical research over the past 20 to 30 years. Generally, this 

research emphasizes cultural burning by American Indian tribes prior to European settlement 

and changes to this fire regime during and after European settlement.

For much of the 20th century, most ecologists did not consider fire to be a factor in the ecology 

of the eastern hardwood biome. Oosting (1942) does not mention fire in his analysis of plant 

communities of the Piedmont region nor does Braun (1950) mention fire in her book on the 

eastern hardwood forests. Whittaker (1956) surmised that fire played a role in hard pine1

 and 

shrub communities in the southern Appalachian Mountains but saw no role for it outside 

these environs. Concomitant with this dismissal of fire as an important ecological process was 

the belief that American Indian tribes had negligible impact on the eastern hardwood biome. 

Indians were believed to be ecologically invisible, living in harmony with nature in a vast forest 

that extended from the Atlantic Ocean to the Great Plains (Adams 1935, Bear 1951, Lillard 

1947, Parkman 1909). However, early writings by European explorers, missionaries, and settlers 

revealed that many American Indian tribes routinely used fire for numerous reasons, especially 

close to their villages and travel paths (Day 1953, Denevan 1992, Hough 1926, Maxwell 1910). 

Unfortunately, historical writings cannot tell us about prehistoric (before written records) fires 

nor do they tell us much about the characteristics of the Indian-set fires.

To help understand prehistoric and Indian-set fires and how they varied by location and time 

in the eastern hardwood biome, we collected and reviewed 30 paleoecology (sediment/soil core) 

and 51 dendrochronology (fire scar) studies. Nearly all of these studies occurred in the past 

20 years and have been published in a wide variety of outlets ranging from master’s theses to 

high-quality, peer-reviewed journals. Because most all these studies are case histories (occur at 

just one site), statistical analysis is descriptive. In this chapter, we look for commonalities and 

trends that provide new insights into the fire regimes of upland oak ecosystems and are useful 

for managers and researchers.

Paleoecology Studies

Paleoecology studies create multi-millennial, coarse-scale chronologies of fire occurrence by 

examining the charcoal that accumulates in undisturbed forest soils and in the sediments of 

small, land-locked lakes. Soil or sediment cores are collected and carefully cut apart in distinct 

layers, and the charcoal fragments are extracted from the layers. The characteristics of the 

charcoal fragments give some indication of the fires that created them. Larger fragments (>2 

mm in surface area) indicate a fire near the sampling site while smaller fragments suggest a fire 

farther away from the study area. Similarly, an abundance of charcoal fragments corresponds 

to a larger, more intense fire while small charcoal accumulations indicate a smaller, less intense 

1

Scientific names of the flora and fauna are listed in the Appendix.
9

fire. The charcoal can also be carbon dated to determine approximately when the fire occurred. 

In some cases, the position of the charcoal in the core can help diagnose the relative time of 

the fire. In the sediment studies, identification and quantification of plant pollen is usually part 

of the project as this indicates the dominant taxa near the research sites and changes in their 

relative abundance through time.

Of the 30 paleoecology studies, 18 were in the central/southern Appalachian Mountains region 

and the remainder was evenly divided between the Great Lakes and Northeast, with none 

occurring in the Midwest (Fig. 9). The chronologies of these studies ranged from 425 to 17,345 

years with most of them producing chronologies of 2,000 to 4,000 years (Table 1). Charcoal 

is found throughout the samples indicating that fire has been a part of the environment for 

millennia. Generally, charcoal was the most abundant in the southern Appalachian sites and 

decreased in abundance to the north. This gradient likely reflects distribution of ancient human 

populations as well as climatic influences. There were more humans and warmer weather in the 

southern Appalachians than there were in the northern Appalachians. Temporally, charcoal 

increased through time, with marked increases at 4,000 years before present (YBP), 1,000 YBP, 

and 250 YBP. The two oldest dates correspond with the rise of Woodland and Mississippian 

Native American cultures and populations in eastern North America while the third marks the 

beginning of European settlement.

Figure 9.—Location of the 

paleoecology (sediment/soil 

charcoal) studies reviewed 

in this project grouped by 

region: Appalachian Mountains 

(AP), Great Lakes (GL), and 

Northeast (NE). See Table 1 

for details of the studies. Note 

the absence of paleoecology 

studies in the Midwest relative 

to the other three regions.

100th Meridian

N

300 miles AP

GL NE
10

Dendrochronology Studies

Dendrochronology studies construct multi-decadal to multi-century, fine-scale chronologies 

from the basal scars that form on living trees when a fire kills a portion of the cambial tissue 

(Fig. 10). Methodology is simple and straight forward for fire scar research. A partial or 

complete cross section is cut from a scarred tree or snag using a chainsaw. The cross section is 

then sanded to reveal the annual rings and the scars (Fig. 11). A calendar year is then assigned 

to the annual rings and scars using standard dating techniques (Speer 2010).

Overall, the 51 fire scar studies were spread throughout the eastern hardwood biome. They 

occurred in 20 states and 1 Canadian province (Fig. 12, Table 2). On average, a study 

consisted of 49 sample trees that produced a 276-year chronology spanning from 1689 to 1965 

(Table 3). In that period, an average of at least 53 fires occurred. This is a conservative estimate 

because it is possible for a fire to burn and not scar trees or have its evidence erased by a later 

fire or decay. The mean fire intervals (MFIs, the time between successive fires) varied through 

time. Prior to European settlement, fires burned about once every 13 years. With settlement, 

fire frequency increased to once every 7 years. After modern fire exclusion policies began, fire 

frequency subsided to once every 46 years. Regardless of the time period, most fires occurred 

during the dormant season when the trees were not actively producing radial growth.

Table 1.—Characteristics of the paleoecology studies reviewed in this project

Regiona State Publication Charcoal type Chronology length

 (years)b

AP GA, KY, NC,TN Welch 1999 Soil not determined

AP NC Delcourt and Delcourt 1997 Sediment 3,900

AP KY Delcourt et al. 1998 Sediment 9,500

AP NC Fesenmyer and Christensen 2010 Soil 4,000

AP TN Cridlebaugh 1984 Sediment 2,800 and 900

AP TN Haas 2008 Sediment 2,800 and 425

AP TN Hart et al. 2008 Soil 6,735

AP VA Kneller and Peteet 1993 Sediment 17,130

AP VA Kneller and Peteet 1999 Sediment 17,345

AP VA Patterson 2006 Sediment 6,000 and 900

AP WV White 2007 Sediment 8,180

GL MN Clark 1990 Sediment 750, 750, and 750

GL NY Clark and Royall 1996b Sediment 10,400

GL ONT Clark and Royall 1995 Sediment 2,000

GL WI Clark and Royall 1996a Sediment 2,400

NE MA Patterson 2006 Sediment 500

NE ME Clark and Royall 1996a Sediment 1,700 and 1,900

NE ME Patterson 2006 Sediment 8,240

NE NY Patterson 2006 Sediment 2,180

NE PA Clark and Royall 1996a Sediment 2,100

aAP = Appalachian Mountains, GL = Great Lakes, and NE = Northeast.

bMultiple chronologies for a publication indicate that the paper reports data from two or more neighboring sites.
11

Figure 10.—A “catface” on the uphill side of a mature 

red pine that probably formed in response to fire 

damage. Photo by Patrick Brose, U.S. Forest Service.

Figure 11. —A sanded cross section of a red pine 

showing the annual rings and dated fire scars. 

Photo by Patrick Brose, U.S. Forest Service.

Figure 12.—Location of the 

dendrochronology (fire scar) 

studies reviewed in this project 

grouped by region: Appalachian 

Mountains (AP), Great Lakes 

(GL), Midwest (MW), and 

Northeast (NE). See Tables 2 

and 3 for details of the studies. 

Note the scarcity of fire scar 

studies in the Northeast relative 

to the other three regions.
12

Table 2.—Characteristics of the fire scar studies reviewed in this project

Regiona State Publication

Recorder 

species

Sample 

size

Chronology 

years

Fire 

years

PreMFIb

(years)

PostMFI

(years)

Mod. 

MFI

(years)

Season

of burnc

AP AL Guyette et al. 2009 Longleaf pine 24 1550-1940 106 2.7 2.6 75.0 DS

AP AL Guyette et al. 2009 Longleaf pine 26 1547-2006 113 3.2 2.5 4.2 DS

AP KY Guyette et al. 2009 Upland pines 31 1742-2005 38 6.3 3.7 85.0 DS

AP MD Shumway et al. 2001 White oak 20 1600-2000 42 8.8 8.0 100.0 DS

AP OH Hutchinson et al. 2008 White oak 137 1870-1984 28 ?d 4.7 24.3 DS

AP TN Armbrister 2002 Upland pines 9 1776-2000 37 7.5 6.7 15.3 n.d.

AP VA Aldrich et al. 2010 Upland pines 63 1704-2003 42 6.0 5.0 30.0 DS

AP VA Hoss et al. 2008 Upland pines 73 1794-2005 53 ? 2.1 4.3 DS

AP VA Lafon et al. 2006 Upland pines 172 1700-2000 227 6.4 3.5 7.4 DS

AP WV Hessl et al. 2011 Upland pines 36 1868-1953 9 ? 8.3 55.0 DS

AP WV Maxwell and Hicks 2010 Upland pines 21 1898-2005 25 ? 3.0 5.0 DS

AP WV Schuler and McClain 2003 Northern red 

oak

17 1846-2002 7 ? 13.3 28.0 n.d.

GL MI Drobyshev et al. 2008 Red pine 248 1707-2006 217 28.5 14.3 20.0 GS

GL MI Guyette et al. 2009 Red pine 30 1494-1914 27 10.2 6.4 ? DS

GL MI Guyette et al. 2009 Red pine 38 1500-2000 19 31.2 11.7 100.0 GS

GL MI Guyette et al. 2009 Red pine 23 1600-2000 11 41.4 17.2 100.0 GS

GL MN Frisell 1973 Red pine ? 1650-1922 32 14.6 6.1 50.0 n.d.

GL MN Heinselman 1973 Red pine 178 1542-1972 71 19.7 3.3 15.0 GS

GL ONT Cwynar 1977 Red pine ? 1696-1960 25 22.8 8.7 50.0 GS

GL ONT Dey and Guyette 1996a White pine 32 1721-1937 14 10.8 12.7 100.0 n.d.

GL ONT Dey and Guyette 1996b White pine 15 1689-1849 9 22.5 ? ? n.d.

GL ONT Dey and Guyette 1996c White pine 18 1634-1875 7 25.2 ? ? n.d.

GL ONT Guyette and Dey 1995a Red pine 26 1665-1780 8 14.0 ? ? n.d.

GL ONT Guyette and Dey 1995b Red pine 18 1636-1994 11 28.3 22.5 64.0 n.d.

GL ONT Guyette et al. 1995 White pine 13 1700-1850 15 8.3 ? ? n.d.

GL WI Guyette et al. 2009 Red pine 28 1706-2005 24 6.5 19.7 75.0 DS

GL WI Guyette et al. 2009 Red pine 15 1705-1870 24 8.6 6.0 ? DS

GL WI Wolf 2004 White oak 16 1804-2000 57 4.5 8.6 4.0 GS

MW AR Engbring et al. 2008 Shortleaf pine 94 1823-2002 61 ? 2.0 2.6 n.d.

MW AR Guyette and Spetich 2003 Shortleaf pine 45 1670-2000 171 11.2 2.8 80.0 DS

MW AR Stambaugh and Guyette 2006 Shortleaf pine 26 1725-2001 92 7.7 2.1 4.0 DS

MW IA Guyette et al. 2009 White oak 20 1699-1860 20 5.0 ? ? DS

MW IL McClain et al. 2010 Post oak 36 1770-1996 118 2.0 1.4 2.5 n.d.

MW IN Guyette et al. 2003 Post oak 27 1654-1999 41 23.0 5.3 7.6 n.d.

MW KS Abrams 1985 Bur oak 19 1862-1983 23 ? 15.7 ? n.d.

MW KY Guyette et al. 2009 Post oak 36 1688-2005 46 9.0 4.2 60.0 DS

MW MO Cutter and Guyette 1994 Post oak 24 1734-1991 45 2.7 8.3 30.0 n.d.

MW MO Guyette and Cutter 1991 Post oak 47 1656-1989 43 4.3 6.4 12.5 DS

MW MO Guyette and Cutter 1997 Shortleaf pine 168 1558-1993 41 15.5 3.7 4.8 DS

MW MO Guyette and Dey 1997 Shortleaf pine 9 1695-1930 67 7.1 2.2 75.0 n.d.

MW MO Guyette and Stambaugh 2000 Shortleaf pine 12 1604-1996 81 5.7 3.7 4.8 DS

MW MO Stambaugh et al. 2006 Bur oak 33 1671-2004 60 6.6 1.6 25.0 DS

continued
13

MW MO Stambaugh et al. 2005 Shortleaf pine 23 1634-1974 97 22.4 1.7 70.0 DS

MW OK Stambaugh et al. 2013 Shortleaf pine 49 1633-1992 88 5.0 1.7 17.0 DS

MW TN Guyette and Stambaugh 2005 Post oak 186 1631-2002 139 8.6 1.2 3.8 n.d.

MW TN Guyette et al. 2009 Shortleaf pine 28 1790-2005 32 6.0 4.5 55.0 DS

MW TX Stambaugh et al. 2011 Post oak 49 1681-2007 30 6.6 6.3 75.0 DS

NE PA Brose et al. 2013a Red pine 93 1530-2010 55 14.3 3.2 100.0 DS

NE PA Lutz 1930 White pine ? 1725-1925 6 37.6 19.5 75.0 n.d.

NE VT Engstrom and Mann 1991 Red pine 20 1815-1987 17 ? 5.1 90.0 n.d.

NE VT Mann et al. 1994 Red pine 32 1504-1851 18 23.1 12.3 100.0 GS

aAP = Appalachian Mountains, GL = Great Lakes, MW = Midwest, NE = Northeast.

bMFI=Mean fire interval: pre-MFI=before European settlement; post-MFI=after European settlement; mod. MFI= modern times since the beginning of forest 

fire control policies.

c

Season of burn: DS = Dormant Season; GS = Growing Season; n.d. = not determined. d? = unknown. 

Table 3.—Summary characteristics of the fire scar studies by region

Region

Sample 

size

Chronology 

years

Fire 

years

Pre-MFIa

(years)

Post-MFI

(years)

Modern 

MFI

(years)

Season of 

burnb

All Regions 49 1689-1965 53 13.1 7 46 DS, 75%

Appalachian 

Mountains

52 1741-1991 61 5.8 5.3 32.4 DS, 88%

Great Lakes 50 1653-1927 36 18.6 11.4 57.8 GS, 50%

Midwest 49 1698-1986 67 8.7 4.4 33.6 DS, 90%

Northeast 48 1644-1943 24 25.0 10.0 91.3 DS, 62%

a MFI=mean fire interval in years: pre-MFI= before European settlement; post-MFI=after European settlement; 

modern MFI= since the beginning of forest fire control policies.

bDS = Dormant Season; GS = Growing Season. Percent indicates the proportion of the fires that occurred in that 

season of burn. 

Table 2.—continued

Regiona State Publication

Recorder 

species

Sample 

size

Chronology 

years

Fire 

years

PreMFIb

(years)

PostMFI

(years)

Mod. 

MFI

(years)

Season

of burnc
14

The number and timing of the fire studies varied by region. The Midwest had the most fire 

history studies (19), and these occurred between 1985 and 2013 (Tables 2 and 3). The Great 

Lakes was next with 16 studies conducted between 1973 and 2009. The Appalachian Mountains 

region had 12 studies that took place between 2001 and 2011. The Northeast had only four 

fire scar studies, but one was the earliest (Lutz 1930) and another was the most recent (Brose et 

al. 2013a). Mean sample size per study was nearly identical among the four regions (48 to 52 

trees per study). The Great Lakes, Midwest, and Northeast had longer chronologies (275 to 300 

years) than the Appalachian Mountains chronology (251 years). This is likely due to differences 

in the longevity and decay rates of the fire-scarred species. The mean number of fire years in a 

chronology was greatest in the Midwest and Appalachians (67 and 61 fires, respectively) than 

in the Great Lakes (36) and the Northeast (24), suggesting a climatic influence. Presettlement 

MFIs were the shortest in the Appalachians (5.8 years), but this may be somewhat skewed by 

a lack of sample size (7) and particularly short MFIs (2.7 and 3.2 years) in northern Alabama. 

Other Appalachian studies produced an average MFI of 7 years. The Midwest had a mean 

presettlement MFI of 8.7 years, and MFIs for the Great Lakes and the Northeast were 18.6 and 

25.0 years, respectively. Generally, transitioning into the postsettlement era decreased MFI by 

about 50 percent. In the Great Lakes, MFI dropped from 18.6 to 11.4 years. In the Midwest, 

the decrease was from 8.7 years (presettlement) to 4.4 (postsettlement). The Northeast had 

the largest drop in MFI, from 25.0 to 10.0 years. The Appalachian Mountains region showed 

no change in MFI. It was 5.8 years before European settlement and 5.3 years afterwards. All 

four regions showed large increases (6- to 9-fold) in MFI after fire exclusion began. In the 

Appalachian Mountains and the Midwest, dormant season fires were the overwhelmingly 

dominant type, but growing season fires composed half of the fires in the Great Lakes and nearly 

40 percent of the fires in the Northeast. These seasonal tendencies did not change through time.

Synthesis

The charcoal and fire scar studies collectively revealed the following findings and trends that are 

informative and useful to forestry managers and researchers:

•	 Fire has been a part of upland ecosystems throughout eastern North America since

the end of the last ice age. Paleoecology studies in the Appalachian Mountains, Great 

Lakes, and Northeast found charcoal dating to at least 5,000 to 10,000 YBP. Fires 

became more common as Native American populations increased about 4,000 YBP and 

again as their cultures developed agriculture about 1,000 YBP. The studies revealed a 

gradient of decreasing fire from south to north and west to east, most likely reflecting 

climatic changes from warm and/or dry to cool and/or moist.

•	 Presettlement (1500 to ~1800 A.D.) fires occurred throughout the eastern oak

biome. We found studies providing pre-1800 data in 18 states and 1 province. 

Additionally, we found references to presettlement fire scars in four other states. Those 

studies and references occurred from Texas to Minnesota and Vermont to Alabama.

•	 Frequency of presettlement fires varied considerably among and within the four

regions of the eastern oak biome. Fire frequencies ranged from annual fires in the 

Midwest to century-long fire-free periods in the Great Lakes and Northeast regions. 

This tremendous variability suggests that lightning was not always the primary 

ignition source, otherwise fire occurrence would have been less variable. Variability 
15

in fire frequency was likely the result of the many American Indian tribes having 

different cultural relationships to fire as well as social upheaval as the tribes came 

into presettlement contact with European influences beginning in the 1500s. The 

introduction of exotic diseases, life-easing inventions (flint and steel), and intertribal 

conflicts over the demand for pelts and skins disrupted American Indian societies 

and led to highly variable fire frequencies. The multi-decadal fire-free periods support 

the contention of Russell (1983) that fires only occurred where and when there were 

American Indian tribes. In the absence of American Indian activity, fire was a rare 

occurrence, especially where climate and topography limited fire spread.

•	 A gradient of decreasing presettlement fire frequency existed from south to north.

Pre-1800 MFIs ranged from 2 to 3 years in Alabama to 30 to 40 years in Michigan and 

Pennsylvania. This gradation likely reflects a climatic influence. There are simply fewer 

periods of time conducive to fire ignition and spread as latitude increases. A less obvious 

gradient occurred from west to east. Generally, MFIs were shorter in the central states 

(Illinois, Missouri, and Oklahoma) than they were in the Appalachian states (Maryland 

and Tennessee), although there were some notable exceptions such as Virginia. This 

gradation likely also reflects climate since droughts are more common and severe in the 

Midwest (Cook et al. 2007), as well as differences in fine fuels (grasses versus leaf litter). 

Both gradients are confirmed by the fire frequency model developed by Guyette et al. (2012).

•	 Presettlement fires generally occurred during the dormant season. Of the 33 studies 

that reported fire seasonality, 26 (78.8 percent) found that the pre-1800 fires burned 

during the dormant season. These dormant season fires may have occurred during the 

autumn for hunting and nut gathering purposes or during the spring to control pests 

and snakes or renew berries and grasses. When growing season fires did occur, the scar 

was situated in the earlywood, suggesting that a dry spring may have extended the 

spring fire season into the early summer. The predominance of dormant season fires 

also indicated that American Indians started these fires because lightning is generally 

restricted to the summer months.

•	 Growing season fires were common in the Great Lakes region and the Northeast.

In these areas, growing season fires typically composed about half or more of the 

presettlement fires. This likely reflects the abundance of pines and lightning as an 

ignition source (Ruffner and Abrams 1998). Growing season fires may have been 

historically important in the xeric pine-oak-heath forests of the Appalachians. Recent 

research at Great Smoky Mountains National Park on unsuppressed lightning fires 

(Cohen et al. 2007) shows that this forest type can ignite from lightning strikes and the 

resultant fires can burn for weeks despite precipitation, eventually covering hundreds of 

acres.

•	 Infrequent prolonged (10 to 40 years or more) fire-free periods are critical to

permitting oak reproduction to grow into the overstory and are a prerequisite to

sustaining upland oak forests. A single extended fire-free period every century or two 

is probably sufficient to sustain the overstory for long-lived (200 to 400 years) species. 

These fire-free periods were common before European settlement (Brose et al. 2013a, 

Guyette et al. 2003). Long fire-free periods in a regime characterized by periodic fire 

favored oak and pine regeneration and set the stage for successful recruitment and 

overstory dominance.
16

•	 Mean fire intervals follow distinct temporal trends. Most of the studies (73 percent) 

showed three distinct fire regimes in terms of MFIs. Collectively, these studies had 

a presettlement MFI of 12.9 years. During European settlement, the overall MFI 

dropped to 6.3 years. Since the advent of fire exclusion policies in the early 1900s, the 

overall MFI has risen to 45.2 years. Most of the studies that did not show this temporal 

trend were in the Appalachian Mountains.

•	 More paleoecology studies are needed. We found no paleoecology studies in the 

literature for the Midwest and several from the Appalachians were lacking complete 

analysis. 

•	 The most suitable species for fire scar research are the hard pines (longleaf, pitch,

red, shortleaf, and Table Mountain) and the white oak group (bur, chestnut,

post, and white). This is likely due to their longevity and ability to compartmentalize 

wounds. The red oak group (black, northern red, and scarlet), other long-lived conifers 

such as eastern white pine and eastern hemlock, and mesophytic hardwoods are of 

lesser value because they are more prone to rot after wounding. The functional time 

span of these species maintaining fire scars seems to be about 500 years as only 2 of the 

51 studies had samples dating to 1500 or earlier.

•	 High-quality fire history studies are lacking for the Northeast region and

the southern Appalachian Mountains. Prior to this project, only three fire scar 

studies existed in the Northeast, and one of those suffered from some sampling and 

methodological shortcomings. In the southern Appalachian Mountains, we found only 

two, but one study was so insufficient in providing data that we could not include it 

in this synthesis. Fortunately, there are several active fire scar studies underway in this 

region that will help elucidate the fire history over the past several centuries.
17

CHAPTER 3: ADAPTATIONS OF OAK TO FIRE

The second part of the fire-oak hypothesis states that oaks1 have adaptations that give them a 

competitive advantage over most other tree species in a periodic fire environment (Abrams 1992, 

Arthur et al. 2012, Brose et al. 2001, Nowacki and Abrams 2008). This chapter reviews some of 

the basic fire and silvics literature to provide an overview of the adaptations that give oaks some 

direct resistance to fire damage during certain life stages. Additionally, this chapter explains how 

oaks indirectly benefit from a regime of periodic fire due to changes in the environment and 

competitive relationships during regeneration and recruitment.

The death of vascular tissue in plants occurs when it is subjected to ≥140 o

F for 60 seconds 

or longer (Hare 1965, Wade and Lundsford 1990, Wright and Bailey 1982). However, 

longer exposure to lower temperatures may also cause the death of tissue. Average maximum 

temperatures from 174 to 700 o

F have been recorded at or near the ground during prescribed 

fires in hardwood forests in the Missouri Ozarks (Dey and Hartman 2005), Ohio (Hutchinson 

et al. 2005a, Iverson et al. 2004, Phillips et al. 2007, Rebbeck et al. 2004), and the southern 

Appalachian Mountains (Elliott and Vose 2005, Greenberg et al. 2012, Phillips et al. 2007). 

These temperatures are capable of causing death to hardwood stems depending on duration 

of exposure and tree characteristics. Fire severity with respect to stem damage and mortality is 

influenced by season, weather, fire type, fire frequency, and factors that affect fire intensity such 

as fuel characteristics (Bond and van Wilgen 1996, Regelbrugge and Smith 1994, Tester 1996, 

Van Wagner 1973). The mortality of seeds and stems following fire is also dependent upon the 

species, location of seed, size of stem, bark thickness, physiological activity, and life stage (Brose 

and Van Lear 2004).

Acorns

Fire effects on acorns can either positively or negatively impact oak regeneration. Fires that 

precede an acorn crop by a few months to a year can promote oak seedling establishment 

(Arthur et al. 2012, Brose et al. 2013b, Wang et al. 2005). Possible mechanisms for this 

promotion include:

•	 Reducing the density and size structure of competing species in the mid and 

understories, thereby increasing light and other resources in the regeneration zone.

•	 Destroying seed of competitors that is stored in the litter.

•	 Reducing the thickness of litter to below 2 inches, thereby eliminating any physical 

barriers to seedling establishment.

•	 Reducing habitat for insects, fungi, and small mammals that feed on acorns.

Conversely, fires that occur shortly after acorn drop cause substantial seed damage and death 

(Fig. 13). Auchmoody and Smith (1993) observed that a low intensity fire killed nearly half of the 

northern red oak acorns in the litter. Greenberg et al. (2012) reported that after a low intensity fire, 

white oak and northern red oak acorns in the litter that were subjected to 400 o

F had high rates of 

mortality, and all acorns perished when temperatures were above 500 o

F. Additionally, surviving 

acorns often produce low-vigor seedlings (Auchmoody and Smith 1993, Dey and Fan 2009).

1

Scientific names of the flora and fauna are listed in the Appendix.
18

Acorns located under matted, wet, and frozen litter are protected from the heat of winter fires. 

Acorns buried in mineral soil have even greater protection from fire damage because soil is a 

poor heat conductor when soil moisture contents are at or below field capacity. For example, 

in low intensity prescribed burns where maximum temperatures reached 325 to 600 o

F just 

above the ground, average soil temperatures were less than 51o

F (Iverson and Hutchinson 2002, 

Iverson et al. 2004). These temperatures did not cause death to buried acorns or roots in the 

soil. Greenberg et al. (2012) found that after a low-intensity dormant season fire, germination 

rates for white oak and northern red oak acorns were lower for seed located on the litter surface 

compared to those in the duff-mineral soil interface or 

buried in 2 inches of mineral soil.

Seedlings

The location of the acorn at the time of germination 

is critical for determining whether an oak seedling 

develops resistance to fire. Acorns have hypogeal 

germination (Burns and Honkala 1990). The cotyledons 

remain in the acorn and this is where the root collar 

and numerous dormant buds form. If the acorn is 

buried in soil by wildlife, then the root collar and the 

accompanying buds are insulated from the damaging 

heat of a fire (Fig. 14). Many of oak’s competitors, such 

as red maple, do not have their seeds buried by wildlife 

plus they have epigeal germination, i.e., the cotyledons 

rise up above the ground to become photosynthesizing 

structures and thus are vulnerable to fire (Fig. 15). This 

difference in mode of germination and location of the 

cotyledons and bud bank confers an advantage to oak 

when fires occur.

Figure 13.—Acorns are one of the life stages 

of oak that are especially vulnerable to fire. 

Photo by Daniel Dey, U.S. Forest Service.

Figure 14.—An oak seedling sprout 

with a well-developed taproot. Note 

the numerous large dormant buds just 

below the root collar. These buds help 

give oak reproduction their tenacious 

sprouting ability. Photo by Daniel Dey, 

U.S. Forest Service.
19

An important physiological trait of oak seedlings that contributes to its survival and persistence 

in a fire environment is that they preferentially store carbohydrates in their root tissue (Fig. 16) 

(Brose 2008, 2011a; Dillaway et al. 2007; Gottschalk 1985, 1987, 1994; Johnson et al. 2009; 

Kolb et al. 1990; Rebbeck et al. 2011). Root carbohydrate level is important for oak seedling 

survival in a periodic fire environment because it supports the growth of sprouts following a fire 

that kills the parent stem. For oak seedlings arising from 

acorns buried below the duff or in soil, the combination 

of an abundance of dormant buds and a reserve of 

carbohydrates stored in root tissue that is protected 

by soil makes them superior sprouters to most of their 

hardwood competitors in a periodic fire environment.

Low- to moderate-intensity surface fires topkill most 

hardwood reproduction, including the oaks (Ferguson 

1961, Lotti et al. 1960, McCarthy and Simms 1935, 

Paulsell 1957, Waldrop et al. 1992). The ability of oak 

seedlings to sprout postfire increases exponentially as 

basal diameter increases (Dey and Hartman 2005). 

Young (e.g., <3 years old) and small (e.g., <0.25-inch 

basal diameter) oak seedlings are susceptible to mortality 

by fire because of their thin bark and relatively low 

root carbohydrate reserves in their small root systems 

(Johnson 1974, Johnson et al. 2009). Larger diameter 

oak seedlings have larger root systems and hence greater 

sprouting capacity (Canadell and Rhoda 1991, Dey and 

Parker 1997a, Johnson et al. 2009).

Figure 15.—Origin of sprouts of white 

oak (WO), red oak (RO), yellow-poplar 

(YP), and red maple (RM). Note that 

the RM and YP sprouts originated 

above ground while those of the oaks 

originated below ground. Photo by 

Patrick Brose, U.S. Forest Service.

Figure 16.—Oak seedlings 

preferentially develop their root 

systems more than their stems. 

This trait helps them survive 

periodic fires and exploit the postfire environment. Photo by Patrick 

Brose, U.S. Forest Service.
20

The key to developing oak reproduction with large root systems is having an adequate amount 

of sunlight in the forest understory (Fig. 17). Sunlight in undisturbed oak stands is often 

<5 percent of full sunlight on mesic and hydric sites, which is below the light compensation 

point for many oak species, and hence is insufficient for net production of biomass (Dey et al. 

2008a, 2008b; Johnson et al. 2009). On these sites, midstory canopies dominated by shade 

tolerant species can develop into overstocked stands, further reducing light levels below that 

needed for oak persistence and dominance. Also, in northern hardwood forests containing 

maple and beech, crown density, depth, and volume is greater because of the shade tolerant 

species compared to pine or oak forests, and this contributes to lower levels of available light in 

forest understories. Removal of the midstory canopy or reducing overstory density is generally 

recommended to provide adequate light for oak root development in forests on better quality 

sites, and the increase in sunlight usually lasts for several years (Brose 2008, 2011a; Brose et al. 

2008; Lorimer et al. 1994; Motsinger et al. 2010; Parrott et al. 2012).

Figure 17.—Dense understory 

shade (top) decreases root 

development of oak seedlings, 

but diffuse shade (bottom) is 

generally sufficient to promote 

development of large oak 

advanced regeneration. Photos by 

Patrick Brose and Daniel Yaussy, 

U.S. Forest Service.
21

Understory light levels on xeric sites are more favorable to promoting oak survival, growth, and 

accumulation of large advance reproduction (Johnson et al. 2009), even without silvicultural 

intervention. Sander (1979) reported and Blizzard et al. (2013) measured average light levels 

of 20 to 25 percent of full sunlight in Missouri Ozark upland oak ecosystems. These forests are 

relatively more xeric than other eastern deciduous forests and lack strong competitors such as 

yellow-poplar that are so prevalent in other ecoregions in the eastern United States. The superior 

drought tolerance and modest nutrient demands of oaks compared to other hardwood species 

also facilitates the accumulation of oak advance reproduction on the more fire-prone, xeric sites 

(Johnson et al. 2009).

Saplings

In general, trees gain resistance to fire damage as they grow larger because increased bark 

thickness improves protection of the cambium. In addition, continued height growth places the 

crown further from the ground, and therefore, the heat generated by surface fires. Bark thickness 

is a major determinant of fire resistance regardless of tree species because bark can insulate the 

cambium and dormant buds from the heat of a fire (Bond and van Wilgen 1996, DeBano et 

al. 1998, Dickmann 1993, Harmon 1984, Hengst and Dawson 1994). Small differences in 

bark thickness produce large differences in fire resistance because the duration of heat required 

to kill tree cambium is proportional to the square of the bark thickness (Hare 1965). Tree 

stem diameter has been correlated to survival after burning because it is directly related to bark 

thickness and tree height, and hence to a tree’s ability to resist heat injury to the cambium or 

crown (Dey and Hartman 2005, Hare 1965, Loomis 1973, Regelbrugge and Smith 1994).

Low- to moderate-intensity fires are effective for eliminating saplings when stem diameters at 

breast height (d.b.h.) are ≤4 inches. (Brose et al. 2013b, Dey and Hartman 2005, Hutchinson 

et al. 2005b). Trees larger than this d.b.h. are seldom topkilled by most prescribed fires, even if 

they are fire sensitive species, because their bark is thick enough to give them resistance to stem 

girdling of the cambium by fire (Fig. 18). Single low-intensity fires often only reduce stand 

basal area by 5 percent (Dey and Fan 2009, Hutchinson et al. 2005b). A single fire usually 

Figure 18.—Single low-intensity fires 

after an extended absence of fire 

will probably have little impact on 

the sapling layer. More intense fires 

or repeat fires are needed to reduce 

the sapling layer and decrease 

understory shade. Photo by Patrick 

Brose, U.S. Forest Service.
22

decreases the density of small saplings (≤3-inch d.b.h.) by 

causing stem dieback and formation of sprout clumps (Henning 

and Dickmann 1996, Paulsell 1957, Thor and Nichols 1973, 

Van Lear and Waldrop 1991, Waldrop et al. 1992). Repeated 

low-intensity burning or a single hot growing season fire can 

increase the amount of basal area reduction (Brose et al. 2013b, 

Fan et al. 2012, Hutchinson et al. 2005b). Furthermore, sprouts 

from former saplings are more susceptible to shoot dieback or 

mortality from additional fires because the stems are smaller in 

diameter. Thus, periodic fires maintain reproduction as seedling 

sprouts or stump sprouts, making them unable to increase their 

fire resistance by growing larger and developing thicker bark. 

In the process of periodic burning, oak reproduction is favored 

(Brose et al. 2013b). Dey and Hartman (2005) and Fan et 

al. (2012) observed differences among species under frequent 

to annual burns for a decade and reported that oaks and 

hickories were superior in persistence compared to most of their 

competitors.

The time since the last fire is a critical variable affecting the 

response of hardwood saplings to periodic fire. A single lowintensity fire after an extended fire-free period (e.g., >20 years) 

often causes little damage to oak’s competitors because they 

have grown large enough in diameter and in bark thickness to 

be fairly fire resistant (Hutchinson et al. 2005b, Waldrop et al. 

2008). Harmon (1984) estimated that red maple could grow 

large enough and have thick enough bark in 23 years to have a 

50 percent chance of surviving a low intensity fire.

Mature Trees

Tree species vary in their ability, as mature individuals, to resist wounding by fire (Fig. 19). 

Several bark properties such as thermal conductivity, specific heat, and thermal diffusivity are 

important in determining its ability to protect the cambium from the heat of fire, and these 

properties vary by species (Hare 1965, Harmon 1984). The most widely recognized bark 

characteristic related to a tree’s resistance to fire injury is bark thickness. Pines are the most fire 

resistance species group largely because of their bark characteristics (Hare 1965, Harmon 1984). 

Among the hardwoods, large (≥20-inch d.b.h.) oaks are generally considered less susceptible to 

fire injury and mortality than most of their competitors because their bark is thicker for a given 

tree size (Harmon 1984, Lorimer 1985). Diameter growth rates are also important because bark 

thickness is primarily dependent on bole diameter. Growth rates vary among species, degree of 

dominance or crown class, and site quality.

Species in the white oak group have thicker bark for given diameters than most other 

hardwoods in the Central Hardwood Region, and they are followed by species in the red oak 

group (Sutherland and Smith 2000). Thinner barked species include American beech, flowering 

Figure 19.—The ability of a tree to withstand a 

surface fire is determined by bark properties and fire 

characteristics which are influenced by a wide variety 

of biotic and environmental factors. Photo by Patrick 

Brose, U.S. Forest Service.
23

dogwood, maples, hickories, and black cherry. Yellow-poplar has thin bark when it is young, 

but it grows rapidly and bark thickness increases such that it is highly resistant to fire injury as 

a large diameter tree (Nelson et al. 1933, Spalt and Reifsnyder 1962, Sutherland and Smith 

2000). Similarly, eastern cottonwood is thin barked and susceptible as a young tree, but it 

grows rapidly and produces thick bark to become one of the most heat resistant species (Hengst 

and Dawson 1994). In contrast, silver maple always has thin bark whether it is small or large 

in diameter (Sutherland and Smith 2000). Large chestnut oaks are highly fire resistant and are 

more likely to survive a high intensity fire than pignut hickory or red maple (Regelbrugge and 

Smith 1994). However, there is substantial variability among the oak species; bur oak is one of 

the most fire resistant in contrast to scarlet oak (Johnson et al. 2009). Protected from fire by 

their bark, oaks are able to live closer to their inherent maximum longevity. Species from the 

white oak group generally live longer than species from the red oak group. For example, post 

oak can live >400 years, whereas scarlet oak seldom exceeds 150 years old. Once oaks make it 

into the overstory as codominant or dominant trees, they can persist for a long time.

Mature overstory trees are also taller, so their crowns are further from the ground making 

them less susceptible to damage from surface fires. A higher intensity fire is needed to kill 

larger (>10-inch d.b.h.) overstory trees (Hutchinson et al. 2005b). However, trying to 

control overstory density with more intense fires is less precise than harvesting trees using the 

shelterwood method, which gives better control over the trees that are removed and the location 

of canopy gaps.

When trees are scarred by fires, they have a suite of defenses that enable them to contain the 

injury and minimize the detrimental effects which include wood decay, loss of structural 

strength with advanced decay, and shortened lives. When oaks are scarred by fires they are 

capable of rapidly and effectively compartmentalizing wounds (Fig. 20) (Smith and Sutherland 

Figure 20.—A cross section of a mature oak from the Missouri Ozarks. Note the numerous 

healed fire scars. This tree has survived many fires through the decades. Photo by Joseph 

Marschall, University of Missouri.
24

1999). This limits the spread of decay throughout the stem and increases a tree’s longevity in 

the overstory where it may potentially be a seed producer for decades. Species in the white oak 

group (bur oak, chestnut oak, post oak, and white oak) are effective at compartmentalizing 

decay (Sutherland and Smith 2000) and are considered resistant to very resistant to decay. 

In contrast, species in the red oak group, hickories, maples, and yellow-poplar are considered 

only slightly resistant or nonresistant (Berry 1969, Nelson et al. 1933, Spalt and Reifsnyder 

1962, U.S. Forest Service 1967). Therefore, large diameter red oak trees are resistant to being 

wounded by low intensity surface fires, but if they receive a large enough wound to permit 

fungal infection, they decay relatively quickly. Trees with decay in their boles are more subject 

to breakage or blowdown in high winds or under the weight of heavy ice and snow loads. 

Wood decay resulting from fire wounds reduces the longevity of the oaks in the overstory by 

increasing the likelihood that they will succumb to stem breakage, blowdown, or death due to 

impaired physiological functioning.

Synthesis

Oaks have a suite of attributes that give them a survival advantage in a periodic fire regime and 

the means to exploit the postfire environment, but their response to fire varies at different stages 

of their life cycle. Some common characteristics of oaks are listed below:

•	 The hypogeal germination trait of acorns results in the root collar and the

accompanying basal buds forming at or beneath the soil surface. This trait protects 

the root collar and buds from the lethal temperatures of a fire. This protection is 

further enhanced if the acorn had been buried by wildlife, which is usually the case.

•	 Oak seedlings emphasize root development in lieu of stem growth. Provided 

there is sufficient light, oak seedlings quickly build large root systems relative to their 

stems. This trait bestows superior survival to oak seedlings relative to their mesophytic 

competitors that emphasize stem height growth more than root development. Once 

the root collar diameter is >0.25 inch, the oak seedling will likely sprout from a 

basal bud if it is topkilled by a fire (Brose and Van Lear 2004). Once the root collar 

diameter is >0.75 inch, the seedling is capable of relatively rapid, sustained height 

growth provided there is sufficient sunlight (Brose 2011a, Brose et al. 2008). The 

emphasis on root development allows oak reproduction to withstand periodic surface 

fires and exploit the postfire environment. Root development also bestows superior 

survival to oaks on dry sites, nutrient poor sites, and during droughts relative to the 

mesophytic hardwoods.

•	 As oaks mature, they develop thicker, more fire resistant bark relative to many

of the mesophytic hardwood species. Bark has several characteristics that convey 

protection from fire. Thickness and texture are two of them. For trees of an equal size, 

oaks will usually have thicker bark than most of the mesophytic hardwood species. 

Additionally, oaks tend to have flaky and/or corky-textured bark that is more resistant 

to heat penetration than solid bark. Among the oaks, those species in the white oak 

group have superior bark for thwarting conductive heat transference than those species 

in the red oak group.
25

•	 When oaks are injured by fire, they are better at compartmentalizing wounds than

most of the mesophytic hardwood species. Because oaks have ring porous radial 

growth and complex cell chemistry (tyloses), they can quickly isolate injuries and 

prevent decay from spreading throughout their boles. This is especially true for species 

of the white oak group. Consequently, oaks can survive injuries longer than many of 

the mesophytic hardwood species, resulting in increasing oak dominance through time.

•	 There are two stages in the oak life cycle that are vulnerable to fire. Fires at the time 

of or shortly after acorn drop are detrimental to oak regeneration because of high acorn 

mortality. Similarly, young oak seedlings are susceptible to mortality, especially from 

growing-season fires, because they have not yet developed large root systems.
26

CHAPTER 4: REVIEW OF FIRE EFFECTS STUDIES

The fourth part of the fire-oak hypothesis asserts that prescribed fire can help regenerate oak 

ecosystems in some situations. To test this assertion, we collected 58 papers produced by 43 

prescribed fire projects conducted in 14 states (Fig. 21) and examined their findings for trends. 

The projects were sorted by stand type (mature, young, or immature), season of burn (dormant 

or growing), and number of fires (single [1], dual [2], or multiple [>2]) based on the site 

descriptions and methods provided in the publications. Stands were considered mature if they 

had a fully stocked overstory, a ubiquitous midstory, and the oak1

 seedlings were either lacking 

or quite small. Generally, these stands had been undisturbed for years or had only recently been 

disturbed by light, thin-from-below treatments. In terms of stand dynamics, mature stands were 

beginning the understory re-initiation stage of development (Oliver and Larson 1990) and the 

oak regeneration process. Young stands included those that were undergoing a shelterwood 

harvest sequence and those that had recently received a final harvest. Oak and mesophytic 

hardwood reproduction was abundant and vigorous, and these stands were near or at the end of 

the oak regeneration process. Also, the time between the harvest and the fire was at least three 

growing seasons. Immature stands were intermediate between young and mature stands. Their 

canopies had recently closed, but they were several decades from being mature. Structurally, 

there were hundreds to thousands of small-diameter trees (3- to 10-inch diameter at breast 

height [d.b.h.]) per acre that were undergoing density-dependent mortality. Immature stands 

were in the stem exclusion stage of stand development (Oliver and Larson 1990).

1

Scientific names of the flora and fauna are listed in the Appendix.

Figure 21.—Map showing the location of the prescribed fire studies used in this project. 

See Table 4 for details of the studies.
27

Dormant season fires occur between leaf abscission in autumn and leaf expansion the following 

spring. During this time, the hardwood reproduction is not photosynthesizing, although sap 

flow may occur in the early spring before leaf expansion begins. Growing season fires take 

place between leaf expansion in the spring and leaf abscission in autumn. The exact starting 

time of the growing season is highly variable and is governed by location, weather, and the 

physiological characteristics of the hardwood species. For example, leaf expansion in mesophytic 

hardwood reproduction usually begins several days to a few weeks before it begins in oak 

reproduction. Therefore, one species group is photosynthesizing while the other is still dormant. 

We considered the beginning of the growing season to be when the leaves of the mesophytic 

hardwood reproduction were ≥50 percent expanded.

After sorting the projects by stand type, season of burn, and number of prescribed fires, we 

examined the results in each paper to determine whether the fire treatment effects on oak 

reproduction were positive, negative, or ambiguous. Positive effects on oak reproduction 

included absolute increases in oak seedling density via establishment of new germinants, relative 

increases to the oak portion of the regeneration pool via differential survival rates between oak 

and mesophytic hardwoods, or acceleration of postfire oak seedling height growth relative to 

that of other species. Negative fire effects on the oak reproduction included decreases in the 

absolute or relative abundance of oak reproduction or loss of the relative oak seedling height 

growth. Ambiguous results were when there was little or no meaningful change in competitive 

relationships between oak and mesophytic hardwood reproduction from preburn to postburn.

Mature Stands

Of the 43 fire projects, 31 (72 percent) took place in mature stands and 29 (93 percent) of 

those involved dormant season fires (Table 4). These were relatively evenly spread among the 

number of fires (11 single-burn, 12 dual-burns, and 8 multiple-burns) and among effects on 

oak (9 ambiguous, 11 negative, 11 positive). However, when these two groupings (number of 

fires and effects on oak) were combined, a pattern of improving benefit to oak as the number of 

fires increased was evident (Table 5). The balance of the first portion of this chapter will review 

some of the noteworthy publications that are representative of the studies conducted in mature 

stands.

Dormant Season Fire

Single Fire

The effect of a single prescribed fire on existing oak seedlings was either negative or ambiguous 

(Tables 4 and 5). Of the 11 studies, 7 found that the number of oak seedlings decreased 

following the fire while the other 4 found no substantial change. Of these, the Johnson (1974) 

study is fairly typical in terms of study design, implementation and outcome.

Johnson’s study took place in southwestern Wisconsin and involved the U.S. Forest Service 

North Central Forest Experiment Station and the Wisconsin Department of Natural Resources. 

The study site was an 8-acre stand dominated by northern red oak. The stand was moderately 

thinned from below (basal area reduced from 120 to 80 square feet per acre) in autumn 1969. 

At this same time, an acorn crop resulted in the establishment of 7,000 new red oak seedlings 

per acre in spring 1970. A year later the stand was split in two, and one section was burned with 
28

Table 4.—Prescribed fire papers reviewed for assessing fire effects on hardwood reproduction organized by stand 

type, season of burn, and number of burns

State Locationa Publicationsb

Stand 

typec

Season 

of burnd

Number of 

burns

Effects on 

oake

AL Bankhead NF Huntley and McGee 1983 M D 1 N

GA Chattahoochee NF Loftis 1990 M D 1 N

IN Purdue Forest Dolan and Parker 2004 M D 1 A

NC Nantahala NF Elliott et al. 2004 M D 1 A

OH Vinton Furnace EF Albrecht and McCarthy 2006 M D 1 A

SC Sumter NF Teuke and Van Lear 1982 M D 1 A

WI Hardies Creek SF Johnson 1974 M D 1 N

WV Geo. Wash. NF Wendel and Smith 1986 M D 1 N

WV Westvaco Forest Collins and Carson 2003 M D 1 N

PA Clear Creek SF Brose et al. 2007 M G 1 N

PA Moshannon SF Gottschalk et al.f M G 1 N

IN Morgan SF Merritt and Pope 1991 M D 2 A

KY Land / Lakes NRA Franklin et al. 2003 M D 2 A

KY Red River Gorge Arthur et al. 1998 M D 2 N

NC Green River WMA Waldrop et al. 2008 M D 2 P

NY Broome County McGee et al. 1995 M D 2 A

OH Zaleski SF Iverson et al. 2008 M D 2 P

OH Zaleski SF Waldrop et al. 2008 M D 2 N

SC Clemson Forest Barnes and Van Lear 1998 M D, G 2 P

SC Clemson Forest Wang et al. 2005 M D 2 P

WI Baxter Hollow Will-Wolf 1991 M D 2 A

WV Fernow EF Schuler et al. 2013 M D 2 P

KY Daniel Boone NF Alexander et al. 2008 M D 3 N

KY Dinsmore Woods Luken and Shea 2000 M D 3 N

KY Red River Gorge Green et al. 2010

Blankenship and Arthur 2006 

Gilbert et al. 2003 

M D 3 A

PA Ft Indiantown Gap Signell et al. 2005 M D 4 P

MO Chilton Creek Tract Fan et al. 2012 

Dey and Hartman 2005 Sasseen and 

Muzika 2004 

M D 4 P

OH Vinton Furnace EF Hutchinson et al. 2012 Hutchinson et 

al. 2005a Hutchinson et al. 2005b 

M D 4 P

MO University Forest Huddle and Pallardy 1996 Paulsell 

1957 

M D 10+ P

TN University Forest Stratton 2007 

DeSelm et al. 1991 

Thor and Nichols 1973

M D 10+ P

AL Bankhead NF Huntley and McGee 1981 Huntley 

and McGee 1983 McGee 1980

Y D 1 N

WI Schmeeckle Reserve Reich et al. 1990 Y D 1 A

CT Goodwin SF Ward and Brose 2004 Y G 1 P

PA State Game Land 29 Brose 2013 Y G 1 P

SC Clemson Forest Geisinger et al. 1989 Y G 1 A

SC Clemson Forest Stottlemyer 2011 Y G 1 P

continued
29

a low-intensity prescribed fire while the other served as an unburned control. Data collected 

that autumn indicated that the burned seedlings had a 40 percent survival rate while the control 

seedlings had a 90 percent survival rate. The fire had killed approximately half of the northern 

red oak seedlings.

One of the criticisms of the Johnson (1974) study is that it reports results collected from one 

inventory conducted just a few months after the fire. A comparable study with a longer interval 

between treatment and inventory is Wendel and Smith (1986). This study occurred in central 

West Virginia and was a cooperative effort by the U.S. Forest Service Northeastern Forest 

Experiment Station, the Washington National Forest, and the West Virginia Department of 

Natural Resources. Like the Johnson study, this site was thinned to 90 square feet per acre basal 

area and burned a year later. Prior to the fire, desirable hardwood reproduction was measured 

at 3,814 stems per acre, and 5 years later the density was 3,500 stems per acre. However, within 

these numbers, the amount of oak dropped by nearly 80 percent while the amount of red maple 

and black locust increased by 17 and 120 percent, respectively. Clearly, the fire had set back the 

oak regeneration process.

Why did these prescribed fires produce such negative results for the oak reproduction? The 

main problem with both of these studies was that the oak seedlings were small and had been 

VA Horsepen WMA Brose 2010 

Brose and Van Lear 1998 Brose and 

Van Lear 2004 Brose et al. 1999a 

Y D, G 1 P

VA Powhatan WMA Keyser et al. 1996 Y G 1 P

PA Allegheny NF Broseg Y G 2 P

WI Jordan Timberlands Kruger and Reich 1997 Y G 2 P

CT Cockaponset SF Ward and Stevens 1989 I G 1 P

NC Duke Forest Maslen 1989 I D 1 A

WV WVU Forest Carvell and Maxey 1969 I D 1 P

aEF = Experimental Forest, NF = National Forest, NRA = National Recreation Area, SF = State Forest, and WMA = Wildlife Management Area. 

bFor studies producing multiple papers, the primary paper is listed first. 

c

M = mature, Y = young, I = immature.

dD = dormant season, G = growing season.

eA = ambiguous, N = negative, P = positive.

f

Unpublished data on file at the U.S. Forest Service Lab, Morgantown, WV.

gUnpublished data on file at the U.S. Forest Service Lab, Irvine, PA.

Table 5.—Distribution of prescribed fire projects conducted in mature oak forests by the 

number of burns and the effect on oak regeneration process. Note the trend of increasingly 

positive effects on oak as the number of fires increase from one to more than two.

----------------------------Number of Fires-----------------------------

Effect on Oak Single (1) Dual (2) Multiple (>2) Total

Positive 0 6 5 11

Ambiguous 4 4 1 9

Negative 7 2 2 11

Total 11 12 8 31

Table 4.—continued

State Locationa Publicationsb

Stand 

Typec

Season 

of burnd

Number of 

burns

Effects on

oake
30

growing in dense understory shade for all of their lives. Consequently, they had small root 

systems with little root carbohydrate reserves and simply could not sprout postfire. Second, the 

prescribed fires were conducted in mid- to late-April so the small seedlings may have already 

begun expanding their leaves, further lowering their root carbohydrate reserves. Finally, neither 

study excluded white-tail deer from the sites, so excessive deer browsing may have subsequently 

eliminated many oaks that sprouted postfire. Regardless of why these studies failed to promote 

oak reproduction, it is evident that prescribed burning can impede the oak regeneration process 

under some circumstances.

Another impact of single prescribed fires on the oak regeneration process is their effect on 

recently fallen acorns. This facet of fire and the oak regeneration process was the earliest one 

reported in the scientific literature. Korstian (1927) found that fires exceeding 400 °F readily 

killed acorns, but there was a mortality gradient among species with acorns of the red oak group 

surviving fire at a higher rate than those of the white oak group. Korstian surmised that this 

gradient was caused by the differences in germination timing (autumn for white oaks, spring 

for red oaks) between the two groups. Subsequent research has confirmed that acorns are easily 

killed by fires (Auchmoody and Smith 1993, Dey and Fan 2009, Greenberg et al. 2012).

Dual Fires

Studies utilizing two prescribed fires showed varying responses (Tables 4 and 5). Most reported 

ambiguous effects (Franklin et al. 2003, McGee et al. 1995, Merritt and Pope 1991) on the oak 

reproduction with some showing positive outcomes (Barnes and Van Lear 1998, Schuler et al. 

2013) and a few showing negative outcomes (Arthur et al. 1998, Luken and Shea 2000). Two 

oak sites that were part of the National Fire and Fire Surrogate Project illustrate this confusion 

as they report differing outcomes between sites as well as among topographic positions (Iverson 

et al. 2008, Waldrop et al. 2008).

The oak sites from the National Fire and Fire Surrogates Project were used to examine the 

responses of hardwood reproduction and many other variables to prescribed fire and mechanical 

fuel reduction treatments in western North Carolina and southern Ohio (Waldrop et al. 2008). 

Dormant season strip-heading fires were conducted twice at both sites with and without a 

mechanical treatment. In North Carolina, the mechanical treatment was chainsaw felling of 

shrubs while overstory thinning was the mechanical treatment used in Ohio. Oak regeneration 

varied by treatment at both sites. In North Carolina, the oaks showed little response to any 

treatment during the first year but increased significantly in number between years 1 and 3 

in the burn only and mechanical plus burn plots. A decrease was observed in year 5 after the 

second burn, but the difference was not significant. The mechanical-only treatment had little 

initial impact on oak regeneration in North Carolina, but a significant increase was observed 

between years 3 and 5. Oak numbers decreased at the Ohio site in all treatment units during 

the first year, although the difference was not significant in the mechanical-only treatment unit. 

No changes occurred between years 1 and 4.

Competitors of oak tended to follow the same patterns in North Carolina and Ohio. Red 

maple showed little response to treatment during the first year in North Carolina, but in 

Ohio there were significant decreases in the number of red maple seedlings in all treatments, 

including the control. Burning with and without mechanical treatment significantly increased 
31

red maple numbers in year 3 (North Carolina) or year 4 (Ohio), but in North Carolina, the 

second burn reduced numbers to pretreatment levels. Yellow-poplar increased over time in the 

mechanical-only plots in Ohio. However, this response was small in comparison to the large 

increase in numbers of yellow-poplar seedlings observed the first year after burning at both sites. 

The numbers of yellow-poplar seedlings decreased by the third measurement at both sites and 

even more after the second burn in North Carolina. This result agrees with the results of Brose 

and Van Lear (1998) who emphasized the need for prescribed burning after yellow-poplar 

seedlings become established. Oaks were 4 to 6 times more numerous after the second burn in 

North Carolina than were seedlings of yellow-poplar. The burn-only treatment changed stand 

structure by reducing the sapling/shrub layer, but it did little to thin the overstory.

Iverson et al. (2008) used the same study area in Ohio as Waldrop et al. (2008) to compare 

treatment impacts as they varied across different positions of the landscape. Study plots in Ohio 

were larger than those in North Carolina (50 acres vs. 20 acres) thus allowing a comparison of 

treatment impacts across dry and mesic sites. The drier landscape positions generally had more 

intense fires, more canopy openness, and more oak and hickory advance regeneration; several 

other tree species also exhibited marked landscape variation in regeneration after treatments. 

Though advance regeneration of several competing species became abundant after the initial 

treatments, the second fires reduced the high densities of the two major competitors, red maple 

and yellow-poplar. The authors suggested that on dry or intermediate sites with at least 2000 

oak and hickory seedlings per acre, reducing canopy cover by 8 to19 percent followed by at least 

two fires should promote oak and hickory to be competitive over about 50 percent of the area. 

However, no appreciable oak and hickory regeneration developed on mesic sites.

The study by Iverson et al. (2008) on relatively large (>50 acre) treatment units showed some 

promise as well as some of the problems that managers face. Though thinning and burning 

increased the density of oak advance regeneration, there also was ample competition from 

species that had different strategies in dealing with the new conditions brought about by 

the thinning and burning. Oak regeneration varied spatially across the large sites because 

topography, fire intensity, and canopy openness were also highly variable. A simple model for 

this region suggests that if a mature forest on a dry or intermediate site with at least 2000 oak 

and hickory seedlings per acre is thinned to 80 percent canopy cover and burned to reduce 

competition and sustain canopy openness, there is a 50 percent chance (or 50 percent of the 

treated area matching these criteria) of oak and hickory successfully becoming competitive for a 

position in the next forest.

Multiple Fires

Even though prescribed burning in hardwoods has been discussed for several decades, it was 

not used on an operational scale until the 1980s (Van Lear and Waldrop 1991). Consequently, 

long-term studies involving multiple fires in hardwood forests are rare, but a number of new 

publications are available describing results after three or four periodic fires. Generally, these 

studies describe positive effects of multiple prescribed fires on oak reproduction, but a few 

report negative effects (Tables 4 and 5).

One of the longest running fire studies in eastern hardwood forests is located in Tennessee 

(Stratton 2007). In 1962, an oak barren in south central Tennessee was divided into 3 
32

treatments (burned annually, burned every 5 years, or not burned) and these treatments 

have been implemented for the past 50 years. Oak has become the dominate species in the 

understory as mesophytic hardwood reproduction has gradually died out, but none of the oak 

reproduction has successfully grown into the canopy in any of the fire treatments. Apparently, a 

5-year fire return interval is too short for oak reproduction to grow large enough to withstand a 

surface fire without being topkilled and forced to sprout again.

Blankenship and Arthur (2006) used a study site on the Cumberland Plateau in eastern 

Kentucky to examine stand structure after prescribed burning two or three times. The same 

study site was used by Green et al. (2010) after another set of burns (three and four burns) to 

examine oak and red maple seedling survival. Burning was conducted by backing fire down 

the ridge and by point source or strip-heading fires if a higher intensity was desired. Fires 

were in the dormant season for the first two fires and then switched to the growing season 

for later burns. Burning altered stand structure by reducing overstory stem density by 30 

percent and midstory stem density by 91 percent (Blankenship and Arthur 2006). Midstory 

oak and red maple stem densities were reduced by 94 and 85 percent, respectively. Damaged 

or dead overstory and midstory stems sprouted, greatly increasing the number of trees in 

the ground layer, with oak, red maple, and dogwood being the most common species after 

three burns. Green et al. (2010) tagged chestnut oak, scarlet oak, and red maple seedlings 

and followed survival and growth through three and four prescribed fires. Burning reduced 

the numbers of seedlings of all three species, but scarlet oak had significantly higher survival 

than chestnut oak and red maple. Scarlet oaks that were burned four times were significantly 

taller than chestnut oak and red maple burned either three or four times. Overall, scarlet oaks 

had better survival and growth than red maples but were not able to eliminate the maples. 

Both papers (Blankenship and Arthur 2006, Green et al. 2010) emphasized that after several 

burns, oak regeneration was in a better competitive position than red maple, but the goal of 

producing predominately oak regeneration had not been reached. Additional burning and/or 

implementation of other silvicultural treatments are necessary to reach that goal.

Alexander et al. (2008), also working in Kentucky, had similar results to Blankenship and 

Arthur (2006) and Green et al. (2010). Numbers of midstory trees were reduced by burning 

one or three times, but sprouting caused large increases in the numbers of trees in the smaller 

size classes. Both single and repeated prescribed burns increased understory light and reduced 

red maple survival. However, neither burning regime placed oaks in an improved competitive 

position. The authors suggest that successful oak regeneration is difficult to predict because 

it is controlled by three highly variable and interdependent factors: life history traits of oaks 

compared to competitors, preburn stature of oak seedlings, and variability of fire characteristics. 

Although not suggested by these authors, other factors that also control oak regeneration are 

understory light levels, site quality, and landscape attributes (aspect, slope, and slope position).

Hutchinson et al. (2005a, 2005b) studied regeneration after two and four dormant season 

prescribed burns on xeric, intermediate, and mesic sites in southern Ohio. Burning, conducted 

by strip-heading fires, had little impact on overstory trees over 10-inch d.b.h. The density of 

smaller trees (4- to 10-inch d.b.h.) was reduced by 31 percent after burning twice and by 19 

percent after burning four times. The two-burn treatment had higher fire intensity, resulting 

in greater mortality of small trees. Burning also reduced sapling density by 86 percent. 
33

Regeneration after burning was numerous and included the same species that were killed by 

burning. In this trial, results were similar among xeric, intermediate, and mesic sites. The 

largest change was brought about by the higher fire intensities associated with the two-burn 

scenario which was better at opening the canopy. In addition, burning at longer intervals 

may have allowed greater buildup of fuels as stems and branches of trees killed by one fire 

fall over and become fuel for the next fire. Waldrop et al. (2010) found that over time, fine 

woody fuels increase in abundance after burning until the next prescribed fire. A factor often 

overlooked is delayed mortality which can occur for several years after a single fire. Yaussy and 

Waldrop (2010) showed that the likelihood of mortality was related to prior tree health, size 

class, species, and first-order fire effects. Hutchinson et al. (2012) concluded that periodic fire, 

coupled with natural gap dynamics, may be a feasible management strategy for perpetuating oak 

forests where harvesting is not an option.

Growing Season Fire

Single Fire

In our survey of the fire and oak literature, we found only three studies reporting results of a 

single growing season fire in a mature stand. The Barnes and Van Lear (1998) study occurred 

on the Clemson University Forest in South Carolina while the Brose et al. (2007) and 

Gottschalk et al. (2013) studies took place in Pennsylvania on the Clear Creek and Moshannon 

State Forests, respectively. Barnes and Van Lear (1998) compared a single growing season fire 

in 1992 to three dormant season fires conducted in 1990, 1992, and 1993. All burns began 

with backing fires and were completed with strip-heading fires. Postburn oak density was not 

significantly different between the two burning treatments. The single growing season burn 

was as effective at promoting open growing conditions as were the three dormant season burns. 

Additionally, burning in the growing season was more effective at reducing competition from 

yellow-poplar. This study suggests that even though burning in the growing season is more 

difficult than in the dormant season, it can be more effective and, ultimately, less expensive.

In the two Pennsylvania studies, postfire sprouting of oak seedlings was 65 percent less than 

that of mesophytic hardwood reproduction. This large difference between the two species 

groups was likely due to the oak seedlings having much smaller root systems relative to the 

larger non-oak reproduction.

Dual and Multiple Fires

None of the mature hardwood studies we reviewed included two or more growing season 

fires. However, a long-term fire study conducted in pine-dominated stands on the Santee 

Experimental Forest of South Carolina does provide some insight into this type of fire regime 

(Langdon 1981, Lewis and Harshbarger 1976, Lotti et al. 1960, McKee 1982). The study was 

established in 1946 with annual and periodic (every 3 to 5 years) burning conducted in summer 

and winter until Hurricane Hugo severely damaged the study in September 1989. Waldrop 

et al. (1992) reported on changes to vegetation through 43 years of treatment. When plots 

were burned every 3 to 5 years, either in summer or winter, trees >5-inch d.b.h. were largely 

unaffected as they were too tall and their bark was too thick to be impacted by low-intensity 

burning. Trees with a d.b.h. between 1 and 5 inches were topkilled gradually over time. These 

stems then sprouted, resulting in a large increase in stems ≤1-inch d.b.h. Annual winter burning 
34

produced similar results, but the largest number of sprouts was in this treatment. With each 

of these treatments, vegetation had at least one growing season to recover from burning. It 

was only in the annual summer burn treatment and only after many burns that hardwoods 

were nearly eliminated from the forest floor. The most resilient species were the oaks which 

persisted through 18 to 20 annual summer fires (Langdon 1981). This result has been cited 

by many authors as an indication of the competitive advantage of oaks over other hardwoods 

in a regime of frequent burning. However, the density of oak competitors increased in all fire 

regimes except annual summer burning, a fire treatment regime that is impractical for most 

land managers.

Young Stands

We found 10 prescribed fire projects conducted in young stands (Tables 4 and 6). Unlike the 

studies in mature stands, only three involved dormant season fires, while eight used growing 

season burns (note that one project provided data on both seasons-of-burn). Similarly, the fire 

studies in young stands were not evenly distributed by number of burns and effects on oak 

reproduction like those conducted in mature stands. Rather, they were concentrated in the 

single fire and positive effects categories.

Dormant Season Fire

Single Fire 

Just three studies reported results of single fires conducted in young stands during the dormant 

season. Huntley and McGee (1981) burned 3-year-old hardwood clearcuts in northern 

Alabama. They found that the dormant season fire reduced the density of yellow-poplar 

reproduction but had virtually no impact on red maple. Density of oak reproduction was also 

unaffected. In central Virginia, Brose and Van Lear (1998) investigated the impact of a single 

dormant season burn on hardwood reproduction in oak shelterwood stands. Like the Alabama 

study, they found decreases in the density of yellow-poplar reproduction, but little reduction of 

red maple density, except where the fires were intense. A follow-up study (Brose 2010) showed 

that these initial findings persisted, especially on the more intensely-burned plots, and stands 

were leading towards eventual oak domination.

Growing Season Fire

Single Fires

Like the dormant season fire studies in young stands, research into the effects of growing season 

fires is sparse. We found eight studies that had growing season prescribed fires as one of the 

Table 6.—Distribution of prescribed fire projects conducted in young oak forests by the 

number of burns and the effect on oak regeneration process. Note the clustering of studies 

reporting positive effects on oak after just one or two burns.

----------------------------Number of Fires-----------------------------

Effect on Oak Single (1) Dual (2) Multiple (>2) Total

Positive 6 2 0 7

Ambiguous 2 0 0 2

Negative 1 0 0 1

Total 9 2 0 11
35

treatments. In central Virginia, Keyser et al. (1996) found that summer fires in oak shelterwood 

stands reduced the density of red maple and yellow-poplar seedlings by 82 and 97 percent, 

respectively, relative to the unburned controls. Oak reproduction decreased by only 11 percent 

following summer burning. Postfire height growth among the species groups was equal. This 

small study spawned a more comprehensive research project that was also conducted in central 

Virginia (Brose and Van Lear 1998, Brose et al. 1999a) and included late spring and summer 

prescribed fires as treatments in oak shelterwood stands. Results from the previous summer 

burn were verified. Densities of red maple and yellow-poplar reproduction declined by 46 and 

72 percent, respectively, while oak seedling density dropped by only 5 percent. Additionally, 

late spring burning resulted in a 45 percent decline in stem density for the two non-oak species 

while oak density actually increased (an artifact of the sampling design). The importance of 

fire intensity was evident in that the largest reductions in stem densities of maples and yellowpoplars occurred where the fires burned the hottest. This relationship between fire intensity and 

oak dominance was still present 11 years later (Brose 2010).

Besides burning in oak shelterwoods, growing season fires after the final harvest have been 

studied to a limited extent. In Connecticut, Ward and Brose (2004) found that mortality 

of black birch ranged from 66 to 86 percent following mid-spring burning in a recently 

regenerated mixed hardwood stand. Mortality of red maple averaged 15 percent but varied 

widely (0 to 100 percent) depending on fire intensity and size of the red maple reproduction 

prior to the fires. Oak mortality averaged just 9 percent with low variability. In Pennsylvania, 

Brose (2013) investigated the effects of mid-spring prescribed fires on hardwood reproduction 

in former oak stands that had recently received the final harvest of a three-cut shelterwood 

sequence. As in Connecticut, black birch exhibited large decreases in stem densities (~90 

percent) while stem density of red maple declined approximately 50 percent. Density of oak 

reproduction was unchanged by the burning, and virtually all oak stems sprouted after the fires. 

Similar to oak, the densities of black cherry, cucumbertree, and serviceberry were the same 

after the fires as before. Besides reducing the stem densities of black birch and red maple, the 

growing season fire equalized the height growth among the various species.

Immature Stands

Only three publications address fire effects in immature stands (Table 4), and all are reviewed 

here. In West Virginia, Carvell and Maxey (1969) studied a sapling stand partly burned by 

an autumn wildfire 5 years earlier. In the unburned portion, yellow-poplar was the dominant 

species in terms of density and size. However in the burned section, oak and hickory 

dominated. They noted that 40 to 70 percent of the saplings that survived the fire had large 

basal scars and concluded that fire was a poor means to manipulate species composition in 

sapling stands given the loss of future timber value.

Maslen (1989) reported on a single high-intensity dormant season strip-heading fire in a mixed 

hardwood pole stand in the Piedmont of North Carolina. This study looked at understory 

characteristics 7 years after burning, giving a slightly longer-term view of fire impacts. By then, 

there were no significant differences in the numbers of oaks and competitors less than 2 feet 

tall because seedlings and sprouts had grown into the next larger size class. In the 2- to 12-foot 

height class, all species including oaks and yellow-poplar were significantly greater in number 
36

than prior to burning. Oaks over 12 feet tall at the time of the fire were essentially unaffected 

by burning and survived the fire. The results of this study indicated that a single prescribed fire 

did little to the stand other than to remove small regeneration and allow sprouts to grow back 

over time.

In Connecticut, Ward and Stephens (1989) reported the long-term (55 year) effects of a 

summer wildfire that burned through part of a 30-year-old mixed hardwood stand in 1932. 

Prior to the wildfire the stand contained approximately 1,050 stems >1.0-inch d.b.h. per acre 

and 74 square feet per acre of basal area. Oak and hickory made up 21.3 and 6.4 percent of the 

stems, respectively, with the balance consisting of birch, maple, and other hardwoods. In the 

years just after the 1932 fire, stem densities and basal area in the burned area dropped by 84 

and 38 percent, respectively, with few differences among species. In the subsequent decades, 

stem densities in the burned area quickly recovered due to sprouting of the fire-killed stems 

before declining due to natural stand thinning. At the same time, basal area gradually increased. 

By 1987, stem densities and basal areas were equivalent in the burned and unburned areas, but 

the burned area contained considerably more oak (160 stems per acre) than the unburned area 

(65 stems per acre). Negative effects of the 1932 fire included the widespread bole damage to 

the trees that survived the fire and the poor stem form of many of the sprouts that developed 

postfire.

Synthesis

Variations in initial starting conditions, study design, and implementation of the prescribed 

fire treatments coupled with the low overall sample size limited the use of traditional statistical 

approaches to analyze the data in Tables 5 and 6. To overcome these methodological challenges, 

we conducted a meta-analysis of the data from 32 of the 42 prescribed fire projects in Table 4. 

Complete results of the meta-analysis are available in Brose et al. (2013b) and are summarized 

below.

In the meta-analysis, we tested five research questions. The first four questions addressed the oak 

regeneration process while the last one examined stand structure. The five questions were: 

1. Does the density of new oak seedlings (germinants) increase postfire?

2. Does oak reproduction basal sprout after prescribed fires at a higher rate than the 

reproduction of mesophytic hardwood species?

3. Does the proportion of oak reproduction increase relative to that of mesophytic 

hardwood species postfire?

4. Does oak reproduction grow as fast as that of mesophytic hardwood species postfire?

5. Does prescribed fire reduce the density of midstory trees of all species?

Meta-analysis results indicated that prescribed fire tended to facilitate the establishment of 

new oak seedlings from acorns, but the trend was not statistically significant (Fig. 22). Oak 

reproduction sprouted at a higher rate postfire than the reproduction of mesophytic hardwoods 

(Fig. 23). However, oak’s superior sprouting ability did not always translate into increased 

relative abundance in the regeneration pool (Fig. 24). In the postfire environment, the height 
37

Figure 22.—The establishment of new oak seedlings (log response ratio ± 95% confidence interval) 

following short-term (≤5 years) and long-term (>5 years) prescribed fire studies. Log response ratios 

significantly above zero indicate an increase in the density of new oak seedlings. Log response ratios 

significantly below zero indicate a decrease in density. Ratios that are not significantly different from 

zero indicate no change in the density of new oak seedlings.

Figure 23.—The relative sprouting (log response ratio ± 95% confidence interval) of released (Rel) 

and suppressed (Sup) oak reproduction in comparison to mesophytic hardwood reproduction following 

dormant season (Dor) and growing season (Gro) prescribed fires conducted throughout the eastern 

United States. Log response ratios significantly above zero indicate that the oak reproduction sprouted 

postfire at a higher rate than the mesophytic reproduction. Log response ratios significantly below zero 

indicate oak reproduction had a lower postfire sprouting rate than mesophytic reproduction. Ratios that 

are not significantly different from zero indicate that the survival rates of the two species groups were 

equivalent.

 

-2

-1

0

1

2

Effect Size (ln R)

≤ 5 Years

> 5 Years

Grand Mean

 

 -3

-2

-1

0

1

2

3

Effect Size (ln R)

Rel_Dor Rel_Gro

Sup_Dor Sup_Gro

Grand Mean

1 Fire

2 Fires >2 Fires
38

Figure 24.—The relative abundance (log response ratio ± 95% confidence interval) of released (Rel) 

and suppressed (Sup) oak reproduction following dormant season (Dor) and growing season (Gro) 

prescribed fires conducted throughout the eastern United States. Log response ratios significantly 

above zero indicate that the proportion of oak reproduction increased in the regeneration pool. Log 

response ratios significantly below zero indicate that the proportion of oak reproduction decreased in 

the regeneration pool. Ratios that are not significantly different from zero indicate that the proportion of 

oak did not change.

-2

-1

0

1

2

Effect Size (ln R)

Rel_Dor Rel_Gro

Sup_Dor Sup_Gro

Grand Mean

1 Fire 2 Fires >2 Fires

growth of oak reproduction was similar to that of many competing mesophytic hardwood 

species (Fig. 25). Within the usual range of conditions under which prescribed fires are 

conducted in eastern oak forests, a fire has limited ability to kill stems (Fig. 26).

After reviewing the fire-oak publications presented in Tables 4, 5, and 6, and the results of 

the meta-analysis, several findings and trends emerged that will be useful to managers of oakdominated ecosystems.

•	 Many factors influence the outcome of a prescribed fire on oak regeneration.

Among these, the important biological factors are the developmental stage of the oak 

stand and the degree of root development of the oak reproduction. Important fire 

factors are season of burn, fire intensity, and their interaction. Finally, critical site 

factors include topography and the disturbance history of the stand.

•	 In mature stands (understory re-initiation stage), as the number of fires increase,

so does the benefit to oak. Single fires and the initial burn of a multi-fire sequence 

provide little, if any, benefit to the oak reproduction and may actually be detrimental in 

the short term. Conversely, multiple burns spread over a decade or more will generally 

benefit the oak component of the regeneration pool via an improved seedbed for oak 

seedling establishment and enhanced understory light conditions for the subsequent 

growth of the new seedlings and any existing oak reproduction.

•	 In young stands (initiation stage), single fires can rapidly benefit the oak

reproduction. This is likely due to differences in root development between the oaks 

and the competing mesophytic hardwoods that give the oaks a higher postfire sprouting 

probability.
39

Figure 25.—The relative height (log response ratio ± 95% confidence interval) of oak reproduction in 

comparison to mesophytic hardwood reproduction following short-term (≤ 5 years) and long-term (> 

5 years) prescribed fire studies conducted throughout the eastern United States. Log response ratios 

significantly above zero indicate that the oak reproduction was taller than the mesophytic reproduction 

postfire. Log response ratios significantly below zero indicate that the oak reproduction was shorter 

than the mesophytic reproduction postfire. Ratios that are not significantly different from zero indicate 

that the heights of the two species groups were equivalent.

 

-1

-0.5

0

0.5

1

Effect Size (ln R)

≤ 5 Years

> 5 Years

Grand Mean

Figure 26.—The reduction of pole and sapling stem density (log response ratio ± 95% confidence 

interval) following prescribed fires conducted throughout the eastern United States. Log response 

ratios significantly below zero indicate that the number of midstory stems decreased while ratios that 

are not significantly different from zero indicate that the postburn densities were unchanged.

-3

-2

-1

0

1

2

3

Effect Size (ln R)

Saplings

Poles

Grand Mean

1 Fire 2 Fires >2 Fires
40

•	 In immature stands (stem exclusion stage), prescribed fire can increase the relative

proportion of oak. However, there will be large economic losses due to bole damage 

to the trees that survive the fire, and due to stem defects (crook and sweep) of the new 

sprouts.

•	 Among the various eastern tree species, postfire sprouting ability of the

reproduction varies widely. Some species are nonsprouters (eastern hemlock and 

white pine), some are poor sprouters (sweet birch and yellow-poplar), some are 

moderate sprouters (red maple), and some are excellent sprouters (oak and hickory). 

The sprouting ability of a species is a function of its capability to form dormant basal 

buds coupled with its germination strategy (epigeal or hypogeal), its juvenile growth 

strategy (root-centric or stem-centric), and its shade tolerance (i.e., the optimal light 

regime for juvenile growth). Sprouting ability is also influenced by season of burn, fire 

intensity, and their interaction.

•	 Growing season fires will have more impact (i.e., kill more stems) than dormant

season fires because the vegetation is physiologically active. Similarly, multiple 

fires kill more stems than single fires due to accumulated damage on midstory and 

overstory trees. In general, a single dormant season fire will have the least impact on 

forest structure while multiple growing season fires will have the most impact. Multiple 

dormant season fires and single growing season fires will have an intermediate impact. 

Within any of these, fire intensity will also play a role as hotter fires have more impact 

than cooler fires.

•	 Immediate mortality from a single fire is typically confined to the regeneration

layer and small saplings (<3-inch d.b.h.). Midstory trees, large saplings (3- to 6-inch 

d.b.h.), and pole-size trees (6- to 10-inch d.b.h.) are periodically killed and some 

may succumb to delayed mortality. Overstory trees (>11-inch d.b.h.) are generally 

unscathed unless there is an accumulation of fuel at or near their bases.

•	 Long-term fire studies in young stands are needed. We found only 10 fire studies 

done in oak shelterwoods or recently-regenerated oaks stands, and just one of 

them reported results more than 10 years postfire. While the vast majority of these 

studies reported positive results for the oak reproduction, more research is needed to 

understand other ramifications of this approach.

•	 Prescribed burning can make the oak regeneration process more difficult. Fire kills 

many of the acorns already on the ground. Small oak seedlings with undeveloped root 

systems are virtually defenseless against a fire, especially a growing season burn. Prescribed 

fires can cause a large influx of new seedlings from seed stored in the forest floor, 

exacerbate invasive species problems, and incite excessive browsing by white tail deer.
41

CHAPTER 5: GUIDELINES FOR USING FIRE IN OAK ECOSYSTEMS

The fire-oak hypothesis is fundamentally sound based on the review of the pertinent literature 

in the preceding three chapters of this report. Fire history research clearly shows that periodic 

fires have been an integral part of the disturbance regime of upland oak1

 ecosystems throughout 

eastern North America for millennia. American Indians were the primary ignition source of 

these fires, but lightning also contributed to fires. Basic silvics research shows that oaks have 

several physical characteristics that give them a survival advantage in a periodic fire environment 

as well as allowing them to exploit the postfire environment. Recent fire research demonstrates 

that prescribed fire can be used in some situations to promote the regeneration of oak. In this 

chapter, we synthesize the findings presented in the previous chapters, especially Chapter 4, into 

guidelines for using prescribed fire to regenerate and restore upland oak ecosystems in eastern 

North America.

Essential Background Information

To correctly use prescribed fire in oak forest management, one must first understand even-aged 

stand development (Fig. 27) (Oliver and Larson 1990), the oak regeneration process (Johnson 

et al. 2009), and how they interrelate (Arthur et al. 2012). The canopies of many eastern forests 

are even-aged or irregular-aged and are dominated by two cohorts of oak resulting from the 

disturbance pattern that occurred widely from the mid-1800s to the early 1900s (Brose et al. 

2001). During those decades, much of the eastern hardwood biome was cleared for agriculture 

or to provide raw materials for the Industrial Revolution and a rapidly-growing nation. In 

many locations, wildfires followed forest clearing, and both of these disturbances ceased rather 

suddenly due to exhaustion of forest resources and conservation efforts. Consequently, the 

burned cutover forests were left to regrow, resulting in young, even-aged hardwood forests 

(stand initiation stage) dominated by oaks. With time, these young forests grew into and 

through the stem exclusion stage and are currently mature stands in the understory re-initiation 

stage. For them to be regenerated to oak–that is to begin the stand initiation stage again–they 

must successfully pass through the regeneration process in a way that favors oaks. Otherwise, 

they will convert to a different forest type.

The oak regeneration process, or the procedure by which mature oaks are replaced by their 

progeny, consists of three phases and an event. The three phases include acorn production, 

establishment of new oak seedlings, and development of the seedlings into competitive-sized 

oak reproduction (Fig. 28). The event is an adequate, timely release (Johnson et al. 2009, 

Loftis 2004). The oak regeneration process usually takes at least a decade due to the sporadic 

occurrence of large acorn crops and the emphasis on root development by young oak seedlings. 

Oaks typically have heavy masting events every 5 to 10 years depending on species and location 

(Burns and Honkala 1990). Acorn crops are subject to numerous environmental factors that 

can slow this phase of the regeneration process or force it to start again (Arthur et al. 2012, 

Johnson et al. 2009). For example, diseases, insects, and weather can ruin acorns before they 

ever fall from the trees or, once on the ground, acorns can be destroyed by these same factors 

or consumed by wildlife. Once an oak seedling cohort forms, these seedlings grow slowly for 

several years as they emphasize root development if adequate resources, especially light, are 

1

Scientific names of the flora and fauna are listed in the Appendix.
42

1

5

4

2

3

Figure 27.—The even-aged stand development model with a two-cut shelterwood sequence. Stages are (1) mature 

stand or understory re-initiation, (2) first removal harvest, (3) final removal harvest, (4) young stand or stand initiation, 

and (5) immature stand or stem exclusion. Generally, oak forests need at least 75 years to complete this cycle. Photos by 

Patrick Brose, U.S. Forest Service.
43

Figure 28.—The oak regeneration process. The three 

phases are (A) acorn production, (B) oak seedling 

establishment, and (C) development of competitive-sized 

oak reproduction. Generally, oak forests need 10 to 25 

years to move through the entire regeneration process. 

Photos by Patrick Brose, U.S. Forest Service.

A

B

C
44

sufficient for oak survival and growth. Like acorn production, the root development phase 

of the regeneration process can be slowed, stalled, or forced to begin again due to numerous 

environmental factors. Preeminent among these are the amount of understory shade, deer 

browsing, and the amount and composition of competing vegetation (Brose 2011a, Miller et 

al. 2004). Eventually, the oak reproduction becomes large enough to successfully compete on 

the site and can be released by harvesting the overstory trees. Because of all these factors, the 

oak regeneration process typically lasts 10 to 25 years (Carvell and Tryon 1961, Clark and Watt 

1971, Sander 1972).

In the even-aged stand development model, the oak regeneration process fits between the 

understory re-initiation stage and the stand initiation stage with some overlap into both stages 

(Fig. 29). The oak regeneration process begins in the understory re-initiation stage when the 

mature oaks produce acorns, some of which become seedlings. Once there is a sufficient density 

of oak seedlings to begin a sequence of regeneration harvests, the parent oak stand begins 

transitioning from the understory re-initiation stage to the stand initiation stage. This transition 

may be abrupt (the overstory is completely removed by one harvest) or gradual (the overstory 

is removed by a shelterwood sequence of two or more partial harvests). Oak shelterwoods 

have elements of both stages of stand development. Overstory trees are still present, but their 

influence on the reproduction diminishes as the degree of cutting increases. The final removal 

harvest marks the end of the oak regeneration process and places the new stand in the initiation 

stage.

The first steps of using prescribed fire in oak forest management are to decide what type of oak 

forest is desired in the future and determine where the prospective stand is in the even-aged 

development model and the oak regeneration process. The desired future oak forest is dictated 

by the management objectives while the current condition is ascertained by an inventory. 

The inventory is an absolute necessity if the management objective is a future oak-dominated 

stand similar in composition to the current one. The inventory can be a simple walk-through 

evaluation by a forester experienced with local conditions, a comprehensive, systematic 

assessment such as those done in conjunction with prescriptive expert systems like SILVAH 

(Brose et al. 2008), or something in between. The inventory must provide basic information 

about the overstory, understory, and regeneration conditions to determine oak regeneration 

potential, as well as to identify potential obstacles to forest renewal and sustainability. After an 

inventory reveals where the stand is at in its development and in the oak regeneration process, 

the correct type of fire can be prescribed and coordinated with other silvicultural practices to 

meet the management objectives.

From our review of the oak-fire literature and the results of our meta-analysis (Brose et al. 

2013b), we have determined three categories of prescribed fire that are appropriate in oak 

ecosystems: (1) seedbed preparation burning, (2) release burning, and (3) ecological restoration 

burning. The rest of this chapter describes these categories of prescribed fire, provides guidelines 

on how to conduct these types of fires, and presents pitfalls that can negate the desired outcome 

of the prescribed burn.
45

1

5

4

2

3

A B

C

Figure 29.—The merger of the even-aged stand development model and the oak regeneration process. Note that acorn 

production (A) and oak seedling establishment (B) occur before the first removal harvest (2), and the final removal harvest (3) 

occurs after the oak reproduction has become large enough to be competitive (C). Photos by Patrick Brose, U.S. Forest Service.
46

Seedbed Preparation Burning

Seedbed preparation burning takes place in mature oak stands that are at the beginning of the 

oak regeneration process. These stands are characterized by three conditions (Fig. 30). First, 

oak reproduction is absent or is so scarce as to be a nonfactor in regeneration planning. In 

addition, the understory contains an abundance of woody non-oak species such as American 

beech, mountain laurel, red maple, striped maple, and witch-hazel that cast a dense shade on 

the forest floor. Finally, thick litter (Oi horizon) and duff (Oe and Oa horizons) layers are present. 

The objective of seedbed preparation burning is to reduce or eliminate the dense shade and 

thick litter so a future acorn crop can find a hospitable seed bed resulting in the formation of an 

oak seedling cohort of sufficient density to begin moving forward through the oak regeneration 

process.

Prescribed fire planning for seedbed preparation burns in oak stands is rather simple and 

straightforward. The proposed burn unit is delineated by natural or man-made boundaries that 

function as the control and contingency lines. Anderson Fuel Model 09 (AFM-09) described 

in Anderson (1982) adequately portrays the fuel bed characteristics and provides reasonable fire 

behavior estimates for the environmental conditions under which most prescribed burns are 

conducted in eastern North America (Brose 2009a). The exception to this is when the oak stand 

has an understory shrub layer of ericaceous (heath) species such as blueberry or mountain laurel 

(Fig. 31). If the heath shrub is deciduous, such as blueberry, then AFM-06 is a good choice for 

planning purposes. Evergreen heath shrubs like mountain laurel are not well characterized by 

Figure 30.—The three characteristics of 

oak stands needing seedbed preparation 

burning are a lack of oak seedlings 

in the regeneration layer (top left), 

an understory dominated by non-oak 

species (top right), and thick duff and litter 

layers (right). Photos by Patrick Brose 

and Robert Long, U.S. Forest Service.
47

any of the 13 AFMs, but AFM-05 is a reasonable representation and AFM-10 and AFM-04 will 

provide a lower and upper bound on what to expect for minimum and maximum fire behavior. 

Generally, strip-head fire is the ignition pattern because the objective is litter and understory 

strata reduction without causing significant damage to the overstory trees. Flame lengths are 

generally <2 feet and rates-of-spread are <5 feet per minute, but heath shrubs can increase these 

parameters to 5 to 10 feet and 5 to 10 feet per minute (Fig. 32). See Waldrop and Goodrick 

(2012) for more details on planning and conducting prescribed fires. Finally, season of burn is 

not strongly important in seedbed preparation burning, although growing season fires tend to 

cause more mortality to the understory than dormant season fires.

Figure 31.—An oak forest with a dense ericaceous (heath) understory of 

blueberry, huckleberry, and mountain laurel. These shrubs increase fuel loadings 

that, in turn, lead to intense fires. Photo by Patrick Brose, U.S. Forest Service.

Figure 32.—A fall prescribed fire in eastern Pennsylvania in an oak stand with a 

dense heath understory. Flame lengths are 5 to 10 feet. Photo by Patrick Brose, 

U.S. Forest Service.
48

Seedbed preparation burning provides several benefits to the oak forest including virtually 

eliminating the litter layer and thinning the duff layer. Generally, fire consumes more than 90 

percent of the litter layer (Brose 2009a; Crosby and Loomis 1974; Loomis 1975, 1977). A thick 

litter layer can be a barrier to oak seedling establishment (Arthur et al. 2012, Johnson et al. 

2009), and blue jays and tree squirrels prefer caching acorns in areas of thin leaf litter (Johnson 

and Adkisson 1986, Steele and Smallwood 2002). The impact of fire on the thickness of the 

duff is less drastic as this layer generally retains some moisture that retards its consumption. 

Still, the duff layer can be decreased by 10 to 20 percent by seedbed preparation burning 

through direct consumption and subsequent accelerated decomposition (Loomis 1975, Van 

Lear and Danielovich 1988). Another major impact of seedbed preparation burning is the 

reduction of the mid- and understory strata and the accompanying dense shade (Fig. 33). Fires 

of moderate intensity (flame lengths ~1 foot) can generally girdle stems with a diameter at 

breast height (d.b.h.) of <3 inches. Removing this tall advanced reproduction and small saplings 

creates an understory light regime of approximately 7 percent sunlight (Miller et al. 2004). 

This low level of sunlight improves oak seedling survival and initiates modest root development 

without eliciting germination en masse from the seed bank or creating a hospitable seed bed for 

light-seeded pioneer species. This level of shade from the overstory can limit the growth of shade 

intolerant reproduction and sprouts of existing advance reproduction that form after the fire.

Other benefits of seedbed preparation burning include recycling the nutrients stored in 

downed woody debris (Blankenship and Arthur 1999, Boerner 2000), stimulating the soil 

ectomycorrhizae community (Stottlemyer 2011), reducing the populations of some acorn 

insect pests (Lombardo and McCarthy 2009, Riccardi et al. 2004, Wright 1986), and xerifying 

the soil surface (Barnes and Van Lear 1998). As fire consumes the woody fuels, essential plant 

growth elements including Ca, K, Mg, and P are released back into the soil and are available for 

Figure 33.—An oak stand in western South Carolina lacking midstory and understory 

layers due to repeated growing season fires. Note the dense vegetation in the 

unburned area at the back of the photo. Photo by David Van Lear, Clemson University.
49

other plants. Sequestered N is volatilized, but soil N increases due to increased soil microbial 

activity. The nutrient needs of new oak germinants are not yet well understood, but increased 

soil nutrient availability reduces the likelihood that low soil nutrient status is a limiting factor 

to their survival and growth. Additionally, fire raises soil pH slightly, further increasing nutrient 

availability. And for a still unclear reason, fire stimulates the growth and reproduction of 

ectomycorrhizae in the soil. These fungi form symbiotic relationships with the fine feeder roots 

of oak seedlings, enhancing the seedling’s ability to absorb nutrients and water from the soil. 

Burning also affects numerous acorn insect pests such as weevils, Valentine moths, and gall 

wasps that attack acorns while they are still in the trees or once they have fallen to the ground. 

These insects spend part of their life cycle on the forest floor, and prescribed fire has been 

shown to reduce their impact on acorns. Finally, seedbed preparation burning xerifies or dries 

the surface layers of the soil so that it is a less hospitable seedbed for the germinants of many 

mesophytic tree species.

Despite the benefits of seedbed preparation burning, there are also drawbacks. First, reduction 

of leaf litter and understory shade can be insufficient and short-lived because the fire intensity 

may be insufficient to kill trees >3-inch d.b.h. and the litter layer can reaccumulate in 2 to 3 

years (Fig. 34) (Crosby and Loomis 1974, Loomis 1975, Stambaugh and Guyette 2006). This 

is especially true for single, low-intensity, dormant season fires. Growing season fires reduce the 

midstory strata quicker and to a greater degree than dormant season fires (Barnes and Van Lear 

1998, Waldrop and Lloyd 1991, Waldrop et al. 1987). Second, timing a seedbed preparation 

burn to shortly precede an acorn crop is impossible. Acorn crops cannot be reliably forecasted 

until mid-summer of the year they are going to occur (Koenig et al. 1994, Perry and Thill 1999, 

Figure 34.—An oak stand in western Pennsylvania treated with one low-intensity, 

dormant season prescribed fire. Note the abundance of dense shade on the forest floor 

and the lack of control of the many sapling- and pole-sized stems. Photo by Patrick 

Brose, U.S. Forest Service.
50

Sharp 1958, Whitehead 1969), so insufficient time for planning or uncooperative weather in 

the season before acorn drop can impact the burn. Alternatively, the fire may be successful 

but the acorn crop may be a failure due to any one of the many factors that can cause acorn 

abortion, resulting in fewer acorns benefitting from the seedbed preparation burning. For these 

reasons, seedbed preparation burning usually involves several fires spread over 10 to 15 years to 

create the open understory, maintain a suitable seedbed, and eventually coincide with an acorn 

crop (Fig. 35). Repeat fires are also more effective in reducing density of competitor sprouts in 

the understory (Brose et al. 2013b). Patience and persistence are necessities when using fire for 

seedbed preparation (Weigel et al. 2012).

Other drawbacks to seedbed preparation burning are the loss of viable acorns and small oak 

seedlings (Auchmoody and Smith 1993, Cain and Shelton 1998). In stands with acorns on 

the ground or a recently-formed oak seedling cohort, refrain from burning and instead use one 

of the individual stem herbicide techniques to reduce understory shading (Brose et al. 2008, 

Kochenderfer et al. 2012). If herbicides are not an option, wait several years before burning 

so that the acorns can germinate and the new oak seedlings have some time to develop their 

root systems, if there is enough light. In deep shade, oak seedlings begin declining once acorn 

reserves are exhausted (Brose 2011a, 2011b; Crow 1992; Dey and Parker 1997b; Loftis 1988; 

Schuler et al. 2013). If this is the case, use dormant season burns to minimize oak seedling 

mortality. Finally, seedbed preparation burning can attract deer that subsequently feed on 

acorns and new oak seedlings and stimulate the establishment and spread of some native and 

nonnative invasive plant species such as hay-scented fern and tree-of-heaven if they are present 

onsite or in adjoining areas (Fig. 36) (Rebbeck 2012).

Figure 35.—An oak stand in southern Ohio that has been treated with four prescribed 

fires over 15 years. Note the diffuse shade and the abundance of vigorous oak 

reproduction. Photo by Daniel Yaussy, U.S. Forest Service.
51

Release Burning

Release burning is a type of prescribed fire used in oak stands that are nearing the end of the 

oak regeneration process to lessen or stop an impending shift in species composition from oak 

to mesophytic hardwoods. Candidate stands have three key characteristics. The stands are on 

intermediate to high quality sites (oak site index50 65 to 80), they are undergoing a shelterwood 

sequence or have just been regenerated via a final harvest (Fig. 37), and they contain an 

abundance of vigorous oak reproduction overtopped by taller competing regeneration of 

mesophytic hardwoods.

Oak reproduction suitable for prescribed burning is typically >1 foot tall with a root collar 

diameter ≥0.5 inch. Adequate oak density can range from several hundred to several thousand 

stems per acre with the larger densities needed on the higher quality sites. Density varies by 

management objectives for future oak stocking at maturity and also depends on the ability to 

do additional release treatments at critical stand developmental stages such as at the onset of 

Figure 36.—An overabundance of deer (left) and invasive plant species (right) can cause problems when using prescribed fire 

to regenerate oak forests. Photos by David Marquis (retired) and Patrick Brose, U.S. Forest Service.

Figure 37. —Shelterwood stands (left) and newly-regenerated stands (right) are appropriate for release burning. Photos by Patrick 

Brose, U.S. Forest Service.
52

stem exclusion. Spatial distribution or stocking of the oak reproduction needs to be widespread 

throughout the stand so that at least 50 percent of the inventory plots contain vigorous oak 

reproduction. The reproduction of the competing mesophytic hardwoods will outnumber and 

overtop the oak reproduction by several thousand stems that are several feet taller, especially 

on the better quality sites. When oak shelterwoods and final harvest stands have the above 

characteristics, they are candidate stands for prescribed burning to release the oak from 

competition as described below.

The Shelterwood-Burn Technique

Correctly implementing the shelterwood-burn technique (Brose et al. 1999a, 1999b) is 

more than simply applying fire to a partially-cut oak stand. The proper application of the 

technique actually begins before the shelterwood harvest while the stand is still uncut or has 

had a low/midstory shade reduction treatment. The first step addresses two questions: (1) is 

there enough oak reproduction at this time to proceed with a regeneration sequence given the 

future oak stocking goal, site quality, and obstacles to stand renewal, and (2) can the stand 

be burned in approximately 5 years? If the first question is answered negatively, then you 

must wait to implement the shelterwood-burn technique until there is an adequate density of 

oak reproduction or institute underplanting to reach the desired density of oak reproduction 

(Dey and Parker 1997a, 1997b; Johnson et al. 2009). Also consider the appropriateness of 

implementing a seedbed preparation burn (see previous section). If the second question is 

answered negatively, then you must make alternative regeneration plans such as using the 

herbicide-shelterwood method which is largely a removal of the midstory and overstory trees 

in the lower crown classes by mechanical or chemical methods to promote the growth of the 

abundant but small oak advance reproduction (Loftis 1990). If both questions are answered 

positively, then proceed with the technique by planning and conducting the first removal cut of 

a two-stage shelterwood sequence.

The purpose of the first removal cut is to create the understory light conditions (30 to 50 

percent of full sunlight) needed to promote rapid root development of the oak reproduction 

while slowing the height growth of the competing mesophytic regeneration (Fig. 38) (Brose 

2011a). Because fire will be used in a few years, planning and conducting this harvest requires 

some extra attention. First, lay out access roads and skid trails so they can double as fire control 

lines in the future. This will expedite the preparation for the prescribed fire, decreasing one of its 

costs. Second, create a 50 percent open canopy by harvesting the low-quality stems, undesirable 

species, and some financially-mature trees (Fig. 39). This is more than a commercial harvest. 

It is necessary to remove unmerchantable overstory trees in the lower crown classes and any 

midstory trees to achieve the 30 to 50 percent of full sunlight. Be sure to cut all unmerchantable 

trees and larger saplings, especially those >3-inch d.b.h., because they have higher probabilities 

of surviving a low intensity fire intact. Cut stems have a high likelihood of sprouting, but the 

sprouts will be susceptible to topkill in a subsequent burn. Alternatively, the midstory and 

noncommercial trees can be stem injected with herbicides (Kochenderfer et al. 2012). In terms 

of basal area, the residual will range from 50 to 80 square feet per acre with the higher residual 

levels being left on the better quality sites. The ideal leave trees are high quality oaks that are 

approximately 15- to 17-inch d.b.h. Trees of this diameter will increase substantially in size and 

value over the next 5 to 10 years, especially if they move from Grade 2 saw logs to Grade 1. It 
53

Figure 38.—Eight-year-old chestnut 

oak seedlings grown in shelterwoods 

of varying residual relative densities. 

Those on the left were in 70 to 90 

percent relative density while those 

on the right were in 20 to 50 percent 

relative density. Photo by Patrick Brose, 

U.S. Forest Service.

Figure 39.—An oak shelterwood in northern Pennsylvania with 50 percent residual basal 

area. This is the ideal structure for developing vigorous oak reproduction that will be 

treated with a release burn in a few years. Photo by Patrick Brose, U.S. Forest Service.
54

is unlikely that there will be enough ideal oaks per acre to meet the recommended residual basal 

area so other trees will have to be kept as leave trees, but be sure not to keep any undesirable 

species that are prolific seed producers such as black birch and yellow-poplar. Finally, manage 

the slash to protect the residual crop trees (Fig. 40). The harvest will create concentrations of 

slash, and such “fuel jackpots” within 10 feet of the bases of residual crop trees can create and 

hold an intense fire that will likely damage or kill them (Brose and Van Lear 1999). Be sure 

the logging contract stipulates that directional felling is used to prevent placing tree tops near 

the bases of residual crop trees or that such slash is removed as part of the harvesting operation 

(Brose 2009b).

The first removal cut is followed by a multi-year waiting period of 4 to 7 years (Fig. 41). This 

period is important for several reasons. First and foremost, this is when the oak reproduction 

develops root systems, a necessary precursor to their ability to sprout postfire (Brose 2008, 

2011a; Brose and Van Lear 2004). The wait also allows the seed bank in the forest floor to 

germinate, at least in part (Schuler et al. 2010). The resultant flush of new reproduction 

probably will contain some seedlings that are potential, long-term competitors to oak such 

as black birch and yellow-poplar. As new seedlings, they are virtually defenseless against a 

prescribed fire. Additionally, this wait allows the fuel bed to develop as the logging slash settles 

and dries and leaf litter accumulates from the residual canopy trees. Finally, the waiting period 

allows the residual crop trees to increase in volume and value, making the final harvest more 

profitable. Leave the stand undisturbed until the oak seedlings have root collar diameters ≥0.50 

inch and are at least 2 feet shorter than the competing mesophytic hardwood reproduction. 

These conditions usually develop within 4 to 7 years, depending on site quality.

The purpose of the prescribed fire is to select for the oak seedlings and against the mesophytic 

hardwood reproduction based on the difference in root development strategies between the 

two species groups. A hot spring burn provides the optimal combination of fire intensity 

and season of burn for maximum benefit to the oak seedlings (Brose 2010, Brose and Van 

Lear 1998, Brose et al. 1999a). Because such a fire will occur in an oak shelterwood, careful 

planning is a must. Use AFM-06 or AFM-10 to represent the fuel loadings of oak shelterwoods 

to predict expected fire behavior (Brose 2009a). Identify the residual crop trees in danger of 

Figure 40.—In the shelterwood-burn technique, slash must be managed so it does not accumulate at the base of residual 

crop trees (left). If it does, the crop tree will be badly damaged or killed by the release burn (right). Photos by Patrick Brose, 

U.S. Forest Service.
55

fire damage due to logging slash close to their bases and take preventative measures to protect 

them (Brose 2010). Strive to burn at the ideal time in the spring season when the mesophytic 

hardwood reproduction has expanded their leaves at least 50 percent but the oak seedlings still 

have closed buds and the overstory is still dormant (Fig. 42). The ideal time to burn varies by 

location and elevation as well as from year to year. For example, the optimal burn window 

generally occurs in late April in southern Ohio but is in mid-May in northern Pennsylvania. An 

Figure 41.—A 4-year-old oak shelterwood in central Virginia ready for a growing season 

release burn. Note the dense regeneration layer and the degree of leaf expansion by 

the understory and overstory trees. Photo by Patrick Brose, U.S. Forest Service.

Figure 42.—The ideal time to conduct a release burn is in mid spring when the oak seedlings are 

still dormant (left) but the competing hardwoods have already expanded their leaves by at least 

50 percent (right). These two photos were taken within 30 feet of each other on May 12, 2011 in 

northern Pennsylvania. Photos by Patrick Brose, U.S. Forest Service.
56

extended winter or an early spring will shift this window backward or forward and will shorten 

or extend its duration. Finally, plan a hot prescribed fire. Flame lengths need to be ≥2 feet 

with rates of spread ranging from 3 to 7 feet per minute (Fig. 43). Although this combination 

of fire intensity and season of burn has consistently produced excellent results in shifting the 

composition of the regeneration pool towards oak, burning outside the hot mid-spring window 

can also benefit oak, but to a lesser degree (Brose 2010, Brose et al. 1999a). Cooler fires and 

those conducted earlier in the spring provide less control of competing mesophytic hardwoods, 

and burns done after leaf expansion of the oak reproduction can reduce their survival and 

decrease their postfire height growth. See Waldrop and Goodrick (2012) for more details on 

planning and conducting prescribed fires.

When done properly, the shelterwood-burn technique will provide several benefits to the oak 

reproduction (Brose 2010, Brose and Van Lear 1998, Brose et al. 1999a). It will kill more 

mesophytic hardwood regeneration than oak reproduction, thereby increasing the relative 

abundance of oak in the advance regeneration pool (Fig. 44). Sprouting oaks will improve in 

stem form and rate of height growth (Fig. 45). Nutrients stored in the leaf litter and slash will 

Figure 43.—A moderate-intensity 

spring fire in central Virginia. Flame 

lengths are 2 to 4 feet. Photo by 

Patrick Brose, U.S. Forest Service.

Figure 44.—The same stand 

shown in Figure 43, but several 

weeks later. The dead saplings are 

yellow-poplar and red maple while 

the green sprouts are oak and 

hickory. The relative abundance of 

oak and hickory increased from 10 

to 70 percent in the regeneration 

pool and this dominance continued 

as the new stand grew into 

saplings. Photo by Patrick Brose, 

U.S. Forest Service.
57

be released back into the forest floor for subsequent use by the sprouting oaks (Blankenship 

and Arthur 1999, Boerner 2000). The ectomycorrhizae in the forest floor will be stimulated 

(Stottlemyer 2011). Berry-producing shrubs such as blueberry, huckleberry, and blackberry 

will be reinvigorated or germinate from seed stored in the forest floor. Besides the food 

benefit to wildlife, blackberry may help the development of oak sprouts by slowing the height 

growth of mesophytic hardwood seedlings and inhibiting the spread of ferns. It also provides 

an alternative supply of browse for deer and hiding cover for oak reproduction, potentially 

reducing deer browsing pressure on oaks.

Based on our collective experience, we see six common mistakes committed by land managers 

implementing the shelterwood-burn technique. They are:

•	 Making the first removal cut before an adequate density of oak reproduction is 

established.

•	 Opening up the canopy less than 50 percent with the first removal cut.

•	 Leaving slash accumulations near the bases of residual crop trees rather than mitigating 

this situation prior to burning.

•	 Burning before the oak reproduction develops sufficient roots and before the 

reproduction is overtopped by competing mesophytic hardwood regeneration.

•	 Burning earlier in the spring than is recommended.

•	 Conducting a low-intensity fire rather than a moderate- to high-intensity fire.

Figure 45.—Prescribed fire can improve stem form and accelerate height growth. The left photo shows 

a single red oak sprout replacing the forked parent stem. The right photo shows the same sprout 15 

years later as a vigorous, well-formed sapling. Photos by Patrick Brose, U.S. Forest Service.
58

Committing any of these mistakes will likely necessitate additional prescribed fires or other 

silvicultural treatments to regenerate oak and avoid undesirable results at the end of the 

regeneration process.

The shelterwood-burn technique does have some drawbacks. The residual crop trees are at risk 

for fire damage and mortality (Brose and Van Lear 1999). This risk is both real and perceived 

because even though a veneer-quality oak is not damaged by the fire, potential buyers may pay 

less money for it because of the threat of staining. Larger red oaks (>11-inch d.b.h.) scarred by 

fire can lose up to 10 percent of the value in the butt log within 15 years of burning (Marschall 

2013). The fire can also kill small oak reproduction that have not yet developed large enough 

root systems necessary for vigorous postfire sprouting (Gottschalk et al. 2013). If native and 

nonnative invasive plant species are in the burn unit, they may expand in coverage or they may 

seed in from adjoining areas (Rebbeck 2012). Deer are attracted to burned areas because the 

sprouting hardwoods are especially palatable and nutritious. In addition, mid-spring prescribed 

fires are probably disruptive to ground-nesting birds such as ruffed grouse, wild turkey, and 

several species of neotropical songbirds and are potentially lethal to herpatofauna just emerging 

from winter hibernation (Beaupre and Douglas 2012).

Postharvest Burning

One approach to mitigating the fire damage risk to crop trees in the shelterwood-burn technique 

is to conduct the final removal harvest before implementing the burn. This alternative is 

called postharvest burning (Brose 2013) and mimics the early 20th century disturbance regime 

that produced many of the current oak stands. If more time is needed for oak seedlings to 

get bigger before burning, the overstory harvest itself may provide a short-term release to the 

oak reproduction by crushing the competing mesophytic hardwood reproduction. Therefore, 

burning may be delayed for 1 to 3 years depending on site quality, but should be done before 

the height of the woody competition exceeds the oak reproduction by >2 feet and average stem 

d.b.h. increases to >3 inches. Postharvest burning has much in common with the shelterwoodburn technique. Both have the same objectives and prerequisites and the timber harvests are the 

same for both approaches. Postharvest burning should be done in mid-spring and strive for the 

same fire intensity as in the shelterwood-burn methods. One place they do differ is in planning. 

Postharvest burning has considerably higher fuel loads of 30 to 40 tons per acre (Fig. 46), so 

AFM-12 should be used in place of AFM-06 or AFM-10 (Brose 2009a). Anticipate flame lengths 

that are >5 feet (10 to 15 feet is not unusual) and a large smoke column, so plan accordingly 

for containment resources and smoke dispersal strategies (Fig. 46). See Waldrop and Goodrick 

(2012) for more details on planning and conducting prescribed fires.

Research on postharvest burning is not common in the scientific literature, but the existing 

publications indicate that the oaks in the regeneration pool benefit from fire at the end of the 

regeneration process (Carvell and Maxey 1969, McGee 1980, Ward and Brose 2004, Ward and 

Stephens 1989). Recent research confirms the findings of these studies (Fig. 47) (Brose 2013).

A somewhat different approach to postharvest burning is the fell-and-burn technique 

(Abercrombie and Sims 1986, Phillips and Abercrombie 1987). This method originated in 

the southern Appalachian Mountains and upper Piedmont regions in the 1980s as a way to 

regenerate or create pine-oak stands on low-quality sites (Fig. 48). As the name suggests, this 
59

Figure 46.—Postharvest fuel loads of oak stands can reach 40 tons per acre (left) and burning them produces high-intensity 

fires and large volumes of smoke (right). Photos by Patrick Brose, U.S. Forest Service.

Figure 47.—A 5-year-old postharvest burn stand 

in northern Pennsylvania. Postharvest burning 

appears to create new, oak-dominated stands 

much like the shelterwood-burn technique. 

Photo by Patrick Brose, U.S. Forest Service.

Figure 48. —A young mixed oak-pine forest in 

western South Carolina created by the fell and 

burn technique. Note the diversity of tree species 

as indicated by their different fall colors. Photo by 

Thomas Waldrop, U.S. Forest Service.
60

technique is a multi-step process. First, all the merchantable trees of the existing stand are 

harvested (Fig. 49). This is generally done in the winter. The following spring when the leaves 

are well to fully developed, all the nonmerchantable stems are felled (Fig. 50). Once their 

foliage is cured and their twigs and small branches dry, the site is broadcast burned (Fig. 51). 

This prescribed burn is conducted during the first summer following harvesting and after the 

hardwood stumps have sprouted, but within 1 to 2 days after a soaking rain (Fig. 52). This 

fire reduces the fuel loading, slows the height growth of oak and other hardwood sprouts, 

Figure 49.—To start creating a mixed oak-pine stand via the fell and burn 

technique, completely harvest all the merchantable trees in the existing stand 

during the dormant season. Photo by Thomas Waldrop, U.S. Forest Service.

Figure 50.—Early in the growing season 

after the commercial harvest, fell all 

remaining hardwoods. Photo by Thomas 

Waldrop, U.S. Forest Service.
61

and prepares the site for the final step of planting the pine seedlings. Planting takes place 

the following winter, and the pines are planted at a fairly wide spacing (15 feet x 15 feet or 

greater). The preferred pine species varies by locale and includes loblolly pine in the Piedmont, 

shortleaf pine in the Ozarks, and pitch and white pines in the Appalachians (Fig. 53). Longterm research of this technique shows that the pine seedlings initially lag behind the hardwood 

sprouts in height growth but become dominant by year 7 to 10 (Waldrop 1997). By year 20, 

this method results in a mixed-species stand dominated by oaks and pines (Fig. 54) (Waldrop 

and Mohr 2012).

Figure 51.—Conduct a growing season prescribed fire several weeks after the 

felling of the noncommercial hardwoods. The several-week wait is essential 

so that the hardwood litter has a chance to dry and the stumps have sprouted. 

Photo by Thomas Waldrop, U.S. Forest Service.

Figure 52. —A fell and burn stand shortly after the prescribed fire. Note the 

unconsumed large diameter boles, branches, and root mat and the general 

openness of the site. It is now ready for planting. The stump in the center of 

the picture will sprout again as will seedlings, seed stored in the soil, and a 

myriad of herbaceous species. Photo by Thomas Waldrop, U.S. Forest Service.
62

Figure 53.—A mixed oak-pine forest in the foothills of the southern 

Appalachian Mountains that was created 10 years earlier with the fell and burn 

technique. Depending on conditions and management objectives, the species 

of pine planted can vary among loblolly, longleaf, pitch, shortleaf, or white. 

Photo by Thomas Waldrop, U.S. Forest Service.

Figure 54.—Generally, the planted pines lag behind the hardwood sprouts for 

the first 10 to 20 years, but they eventually catch the oaks in height growth 

and form a mixed oak-pine ecosystem. Photo by Thomas Waldrop, U.S. 

Forest Service.
63

Ecological Restoration Burning

The previous two categories of prescribed burning (seedbed preparation and release) had as 

underlying assumptions timber production as the primary management objective and complete 

harvesting of the mature stand during the course of the regeneration process. In ecological 

restoration burning, timber harvesting is of secondary importance, and the driving objective is 

the reintroduction of periodic fire as an ecological process to restore or rehabilitate the structure 

and function of woodland and savanna ecosystems. Often, the foci of this type of prescribed fire 

are the creation and maintenance of a somewhat open stand structure and a diverse understory 

plant community. Fire is a critical part of the process necessary to sustain woodlands and 

savannas, for without it, these ecosystems would rapidly develop forest structure and function 

in eastern North America where precipitation is adequate and soils are sufficiently productive 

to support tree growth (Grimm 1983, Nuzzo 1986). Implicit within this objective is the 

development of a sufficient quantity of vigorous oak reproduction to replace the mature oaks 

when they succumb to natural mortality events. Ecological restoration burning is rather broad 

and inclusive but is well exemplified by prescribed fires used to create and maintain open oak 

woodlands, savannas, and shrubland habitats.

Open Oak Woodlands

Early European explorers and settlers frequently described parts of the eastern oak forest as 

open and park-like, consisting of large, widely-spaced trees with a diverse understory plant 

community (Fig. 55) (Bromley 1935, Day 1953, Maxwell 1910). While the amount and 

distribution of open oak woodlands is unknown, periodic fire is the most logical ecological 

process responsible for the creation and maintenance of this now rare habitat type. Oak 

woodland restoration burning is almost identical to seedbed preparation burning. Initially, both 

methods strive to decrease the stem density of the midstory strata, thereby gently increasing 

ambient light levels on the forest floor. Both reduce the thickness of the forest floor, creating a 

favorable seedbed for acorns, and promoting the development of grasses and forbs characteristic 

of woodlands. Both similarly impact the acorn insect pests that inhabit the forest floor as well as 

recycle nutrients sequestered in the fine woody debris.

Figure 55.—An open oak woodland 

created by repeated prescribed fire. 

Note the diffuse shade and diverse 

herbaceous layer; two conditions 

sometimes described by early 

European settlers. Photo by Patrick 

Brose, U.S. Forest Service.
64

Conceptually, there are several important differences between seedbed preparation burning 

and ecological burning for woodland restoration. First, the sizes of the burn units usually 

differ. Seedbed preparation burns are done at the stand level or smaller scale by hand (drip 

torch) ignition while woodland restoration burns are generally implemented at a multi-stand 

to landscape scale by aerial (helicopter) ignition (Fig. 56). Second, protection of mature oak 

trees differs. In seedbed preparation burns, protecting mature trees is a high priority because 

they are the seed source for future seedlings and will be harvested once stocking goals for oak 

reproduction are satisfied. In woodland restoration burning, tree protection is much less critical 

(Fig. 57). It is still important for future acorn production, but scarring of mature trees is much 

less serious because these trees will likely never be harvested or their cutting will be to meet 

management objectives other than timber production. Once there is sufficient oak reproduction 

Figure 56.—Prescribed burning to restore open oak woodlands is often 

done at the landscape scale (top) using aerial ignition (bottom). Photos 

by Michael Bowden, Ohio Division of Forestry, and Thomas Waldrop, U.S. 

Forest Service.
65

to meet stocking goals, the two prescribed fire types differ in the implementation of future 

burns. At this point in seedbed preparation, fire is discontinued. No more burning is done as 

the stand passes through the oak regeneration process unless necessary, i.e., release burning 

to control mesophytic hardwood competition. In woodland restoration burning, prescribed 

fire will continue on a periodic basis. The exact periodicity of future burns depends on local 

conditions and management objectives and can be quite variable. As established in Chapter 2, 

the historic mean fire intervals (MFIs) varied among and within regions. Periods of frequent fire 

were common as were multi-year and multi-decadal fire-free periods. This variability in MFIs is 

a critical component in oak communities where fire is an important ecological process. Frequent 

fires are essential when fire has been excluded for too long and to help develop woodland flora 

and reduced the shrub and tree cover in the mid and understory layers. However, sufficient oak 

advance reproduction is needed when preparing to replace the overstory, and this may require 

the establishment of new oak seedlings when sufficient numbers are lacking to meet future 

overstory stocking goals. Sufficiently long fire-free periods are crucial for the recruitment of the 

oak regeneration into larger size classes so trees are large enough to withstand future fires and 

are permitted to continue to growth into the overstory.

Planning for ecological restoration burns in oak woodlands is rather simple and straightforward. 

The proposed burn unit is delineated by natural and/or man-made boundaries that function 

as the control and contingency lines. Anderson Fuel Model 09 (AFM-09) adequately portrays 

the fuel bed characteristics and provides reasonable fire behavior estimates for the vast majority 

of environmental conditions under which most prescribed burns are conducted in eastern 

North America (Brose 2009a). The exception to this is when the burn unit has an understory 

shrub layer of ericaceous (heath) species such as blueberry or mountain laurel. If the heath 

shrub is a deciduous species such as blueberry, AFM-06 is a good choice for planning purposes. 

Evergreen heath shrubs like mountain laurel are not well characterized by any of the 13 AFMs, 

but AFM-05 is a reasonable representation and AFM-10 and AFM-04 will provide a lower and 

upper bound on what to expect for minimum and maximum fire behavior. Because woodland 

restoration burning is generally done on a multi-stand basis using aerial ignition, spot-head 

firing often is the ignition pattern used because this method creates a mosaic of fire intensities 

from creeping, low-intensity backing fires to running, high-intensity head fires. See Waldrop 

and Goodrick (2012) for more details on planning and conducting prescribed fires.

Figure 57.—In forests managed for 

timber production, fire damage like 

that pictured here is unacceptable. 

However in woodland restoration 

burning, bole damage is much less 

critical. Photo by Patrick Brose, 

U.S. Forest Service.
66

Season of burn is very important in woodland restoration burning because other means to 

reduce midstory stem density are often not employed or are unavailable. Growing season fires 

should be emphasized at the beginning of restoration because they will cause more mortality to 

the understory strata than dormant season fires. Once the desired stand structure is achieved, 

dormant season fires can be used to maintain it. Reduction of overstory density to create open 

woodlands can be achieved by increasing fire intensity, stem injection with herbicides, or by 

timber harvesting.

The drawbacks of woodland restoration burning and the mistakes made when implementing 

it are the same ones detailed for seedbed preparation burning. The most common mistakes are 

generally made at the beginning of restoration and include not using growing season fires and 

not burning frequently enough. Both of these result in little substantive change in midstory 

stem density. Another mistake is using fire too frequently once restoration is achieved, and 

regeneration to replace the overstory is needed. This prevents successful recruitment of the oak 

reproduction into large size classes so canopy tree replacement can eventually occur.

Oak Savannas

Savannas are characterized by grass cover with widely-spaced trees (Fig. 58) (Anderson et al. 

1999, McPherson 1997, Nelson 2010). The ground flora is a mixture of grasses and forbs 

common to prairies and woodlands. Scattered patches of shrubs may occur on more fireprotected sites. Tree cover may vary from 5 to 50 percent but must be low enough to permit 

dominance by grasses and other sun-loving plants. Historically, savannas occurred along 

the edges of prairies, representing a compositional and structural transition from prairie to 

woodland and forest. Savannas were prevalent on flat to gently rolling lands of low topographic 

roughness where fires burned more frequently in the past and retarded the encroachment of 

trees and shrubs (Batek et al. 1999, Grimm 1983, Stambaugh and Guyette 2008).

Figure 58.—Oak savannas are characterized by widely-spaced trees and a diverse 

herbaceous community of forbs and grasses. In eastern North America, oak 

savannas only occupy a small fraction of their historic range. Photo by Daniel Dey, 

U.S. Forest Service.
67

In North America, savannas were a major terrestrial biome before widespread European 

settlement. Nuzzo (1986) estimated that about 30 million acres of oak savannas occurred 

throughout the central United States in 1800, but today <0.02 percent of the original 

savannas remain. The greatest loss of the original savannas occurred when they were converted 

to agriculture production. Those that remained rapidly succeeded to forests when fire was 

suppressed (Fig. 59).

In areas of intermediate rainfall (25 to 50 inches per year), savannas are ecologically unstable 

and must be maintained by frequent fire (Mayer and Khalyani 2011, Starver et al. 2011). 

Despite relatively low mature tree density in oak savannas, an abundance of oak seedling 

sprouts persist in the ground layer of vegetation, and when a sufficient fire-free period occurs, 

these sprouts grow rapidly and form closed tree canopies in about 20 years (Cottam 1949, 

Curtis 1959, Nelson 2010). If they are allowed to grow large enough, the trees gain resistance 

to being killed by subsequent fire.

Restoration of oak savannas entails reducing the density of trees and reintroducing fire (Fig. 

60). Most understories of overgrown savannas have 3 to 5 percent of full sunlight, and much 

more light is needed to stimulate germination, promote growth, and encourage seed production 

of the herbaceous species commonly found in savannas. For example, for grasses to proliferate, 

tree canopy cover needs to be <50 percent (Anderson et al. 1999, Mayer and Khalyani 2011, 

Nelson 2010, Starver et al. 2011). Managing overstory crown cover between 5 to 50 percent 

influences the development of the ground flora. Tree crown cover below 30 percent is needed 

to promote grass domination and proliferation of sun-loving forbs commonly found in prairies 

(Nelson 2010). Higher tree cover begins to favor the abundance and diversity of more shade 

tolerant forbs characteristic of woodlands as the grasses decrease in dominance. Maintaining 

tree cover above 50 percent inhibits dominance by graminoids in the ground layer.

Figure 59.—The extended absence of fire allows oak savannas to succeed to 

closed-canopy forests. Once they have reached this point, they are difficult to restore. 

Photo by Daniel Dey, U.S. Forest Service.
68

Law et al. (1994) published a chart useful for managing stand density to produce desired crown 

cover in upland oak savannas depending on overstory tree size and density. For example, if the 

average d.b.h. of overstory savanna trees is 20 inches, then 17 open-grown trees per acre will 

provide 50 percent crown cover. If the desired tree crown cover is only 30 percent, then 11 trees 

per acre that average 20 inches d.b.h. will provide the desired crown cover if the trees are open 

grown.

Fire by itself is a mediocre means to reduce basal area and stocking and increase understory 

light (Fig. 61). In a closed-canopy forest, low-intensity surface fires only increase understory 

light up to about 15 percent of full sunlight by reducing the sapling layer (Lorimer et al. 1994, 

Motsinger et al. 2010, Ostrom and Loewenstein 2006). Moderate to high intensity fires, 

especially in the late spring and summer, are capable of killing larger trees as well as increasing 

the proportion of smaller stems that do not sprout postfire (Brose and Van Lear 1998, Brose 

et al. 2013b). Killing larger, older trees often leads to complete mortality because they are less 

likely to sprout after death of the stem than are saplings and pole-sized trees (Dey et al. 1996). 

Figure 60.—Combining timber harvesting (left) with repeat prescribed burning (right) is the fastest way to restore an oak 

savanna that has deteriorated due to the extended absence of fire. Photos by Daniel Dey, U.S. Forest Service.

Figure 61.—By itself, prescribed 

fire will not restore an overcrowded 

oak savanna. The extended 

absence of fire has changed fuel 

loadings and properties and has 

allowed too many trees to grow 

large enough to withstand fire. 

Note the abundance of midstory 

trees and accompanying dense 

shade despite several prescribed 

burns in the past decade. Photo by 

Daniel Dey, U.S. Forest Service.
69

This can be accomplished at a fine, local scale by placing downed, cured tree tops at the bases 

of live trees or by broadcast burning when fuel moisture and relative humidity are low and fuels 

are sufficient to generate high fire temperatures for the duration needed to kill cambial tissue. 

Managing the reduction of larger trees with fire can be problematic because burning under 

weather and fuel conditions needed to achieve high-intensity fires is risky from a fire escape and 

safety standpoint, and fire is indiscriminant in what trees are killed. Generally, other practices 

such as timber harvesting or chemical/mechanical thinning must be employed with fire to 

complete the restoration process, especially when fire or other major disturbances have been 

absent for decades (Fig. 62).

Timber harvesting to reduce the density of large trees is an efficient alternative method of large 

tree removal and affords a high degree of control of the spatial arrangement and composition 

of remaining trees. It also provides needed income to pay for other costs of restoration. 

Mechanical cutting of large diameter trees before burning is an effective way of controlling 

overstory density. Although larger diameter trees are less likely to sprout, they may produce 

sprouts after mechanical cutting, however, their sprouts are then vulnerable to being topkilled 

by a subsequent fire due to their small diameter and thin bark. If large trees are unmerchantable 

or timber harvesting is not a management option, another effective method is the stem 

injection of herbicides to reduce tree density and cover of larger trees. When applied correctly, 

herbicides can kill trees and their root systems completely, preventing them from sprouting. 

Dead trees can be left standing to fall apart over time, if public safety is not an issue. In the 

process of decaying, snags can provide critical habitat for a wide variety of wildlife. Methods 

of mechanically girdling trees are also available for killing larger trees, though they are not as 

effective as herbicides.

In most cases, reducing tree density in combination with fire produces the fastest improvements 

in ground flora diversity and dominance on sites not previously degraded by livestock 

overgrazing and soil erosion. Prescribed burning alone generally produces increases in 

herbaceous species coverage and richness when trying to restore oak savannas from mature 

forests conditions, but improvements are most dramatic where the overstory is opened up either 

by the fire or companion thinning/harvesting operations (Hutchinson 2006, Hutchinson et al. 

Figure 62.—Long-term management of oak savannas has to include occasional fire-free periods of sufficient duration so 

oak reproduction (left) can gain the necessary size to withstand the surface fires and eventually replace the existing canopy 

trees (right). Photos by Daniel Dey, U.S. Forest Service.
70

2005b, Kinkead et al. 2013, Waldrop et al. 2008). Even without fire, positive short-term responses 

in herbaceous richness and coverage have been achieved by timber harvesting (Kinkead et al. 

2013, Zenner et al. 2006). Hardwood and shrub regrowth after stand thinning without fire 

diminishes gains in herbaceous flora in a few years as shade from woody canopies develop.

In addition to using fire to restore and maintain the woody structure of savanna ecosystems, 

the restoration of fire as a disturbance that shapes the ground flora community is important. 

Fire promotes germination and growth of herbaceous plants by: removing litter that acts as a 

physical barrier to germination and seedling establishment; preparing a receptive seedbed for 

colonization by wind and animal dispersed seed; breaking chemical and thermal seed dormancy 

by producing heat and smoke; releasing nutrients tied up in the litter; and increasing light 

and temperature in the regeneration zone by removing litter and reducing woody structure, 

which stimulates germination and growth. Fire can be managed to favor dominance of 

grasses, legumes, or forbs by varying the season, intensity, and frequency (every 1 to 3 years) 

of prescribed burning (Anderson et al. 1999, Nelson 2010). Simple recommendations cannot 

be given here because vegetation response to fire is complicated by differences in climate, soils, 

hydrology, grazing, and overstory interactions. However, spring fires favor grass domination 

in general while summer burns favor forb diversity (Nelson 2010). Dormant season fires have 

the least impact on the herbaceous plant community (Hutchinson 2006). Grasses thrive under 

annual to biennial burns, but shrubs, trees, and vines increase in dominance with longer fire-free 

periods. Consistent application of prescribed fire tends to create homogeneity in the vegetation 

community, therefore it is recommended to vary the frequency, intensity, and season of burning 

when maintaining savanna ecosystems (Nelson 2010).

Prescribed fire planning in oak savanna restoration and maintenance is rather simple and 

straight-forward. The proposed burn unit is delineated by natural or man-made boundaries that 

function as the control and contingency lines. At the beginning of the restoration process, if 

there has been no mechanical reductions of basal area and stocking, Anderson Fuel Model 09 

(AFM-09) adequately portrays the fuel bed characteristics and provides reasonable fire behavior 

estimates for the environmental conditions under which most prescribed burns are conducted 

in eastern North America (Brose 2009a). Generally, spot-head fires lit via aerial ignition are 

the firing pattern because the objective is litter and understory strata reduction without causing 

significant damage to the overstory trees. Flame lengths are generally <2 feet and rates of spread 

are <5 feet per minute. If the overstory has been reduced by harvesting, then use AFM-06 or 

AFM-10 to represent the fuel loadings to predict expected fire behavior (Brose 2009a). In 

restored oak savannas, the grasses and other herbaceous plants strongly influence fire behavior. 

These are best represented by the grassland fuel models, especially AFM-03 (Grabner et al. 1997, 

Grabner and Dwyer 2001). See Waldrop and Goodrick (2012) for more details on planning 

and conducting prescribed fires.

Typically restoration of savannas takes place on a larger landscape, at least on public lands like 

the Mark Twain National Forest (U.S. Forest Service 2005). The replacement of overstory trees 

on large-scale restoration projects does not require that the entire area forgo prescribed burning 

when recruiting to replace overstory trees. Given that overstory trees may live for 150 to over 

400 years and that it only takes a relatively few trees per acre to provide the desired tree crown 

cover in savannas, infrequent recruitment on smaller acreages can be sufficient to maintain 
71

savanna overstories. By continually permitting recruitment to occur on additional smaller 

areas, a larger savanna landscape can be sustained in perpetuity. The minimum area needed for 

recruitment depends on the size of the overall landscape and the length of the management 

rotation, or average longevity of the overstory trees (i.e., 100, 200 or 300 years).

Eventually, overstory trees need to be replaced because individual trees do not live forever. Post 

oak is one of the longer lived oak species and individuals may live to be over 400 years old, but 

even they need to be replaced to sustain oak savannas. Individual tree longevity is shortened when 

the boles of trees are scarred by fires and wood decay develops, structurally weakening the trees. 

Recruitment of seedling sprouts and grubs into the overstory requires a sufficiently long fire-free 

period for trees to grow large enough in size to gain resistance to being topkilled by the next fire. 

In general, this may require a 20 to 30 year fire-free period depending on tree growth rates and 

source of reproduction (Johnson et al. 2009, Wakeling et al. 2011). Oak stump sprouts are the 

fastest growing source of oak reproduction. One of the fastest growing oak species in the uplands 

is scarlet oak, and its stump sprouts can achieve diameters averaging 3 inches in 10 years when 

growing in the open (Dey et al. 2008b). White oak, bur oak, and post oak are slower growing 

species. Increasing overstory density reduces the growth of oak sprouts and lengthens the time 

needed for them to achieve the bark thickness to become resistant to topkill by fire. These longer 

fire-free periods are not atypical in fire history records for the period before European settlement 

(Guyette and Spetich 2003, Guyette et al. 2002, Stambaugh et al. 2006).

Oak Shrublands

These rare habitats occur in distinct locations due to edaphic limitations, often excessivelydrained, nutrient-poor soils, and are dominated by shrubs and small trees. One such ecosystem 

is the scrub oak ecosystem of the central and northern Appalachian Mountains that is found on 

rocky ridgetops and other xeric environments (Fig. 63). The principal oak species is bear oak, 

a deciduous shrub or small tree. Mountain laurel, huckleberry, blueberry, and sweet-fern are 

common associate shrubs. Depending on the degree of edaphic limitation and the time since 

Figure 63.—Scrub oak 

communities are ecological 

rarities—a result of edaphic 

conditions and periodic fires—

that have tremendous biodiversity 

value. Photo by Patrick Brose, 

U.S. Forest Service.
72

the last disturbance, stunted chestnut oak, scarlet oak, and pitch pine may be present in limited 

numbers. This shrub ecosystem is valued for its uniqueness, scarcity, and as wildlife habitat. 

Of the shrubs, bear oak is especially important because it is the food source for the caterpillars 

of several rare butterflies and moths as well as a source of hard mast for other wildlife species 

(Grand and Mello 2004).

Periodic prescribed burning of the oak scrub community is necessary to renew and regenerate 

the shrubs as they become decadent over time. Fire also prevents the tree species from 

becoming too large and numerous, thus causing shading and nutrient competition problems 

for the shrubs (Fig. 64). Depending on the amount of encroachment by tree species, initial 

restoration burns may be as frequent as 3 to 5 times a decade. Once tree encroachment is 

controlled, fire frequency can be reduced to 1 or 2 fires a decade. Growing season burns are 

generally recommended for initial restoration, but they must be employed with caution in this 

shrub community. Mountain laurel and deciduous heath shrubs are highly flammable at the 

time of leaf expansion because of volatile oils and resins. Flame lengths that are >20 feet are 

not uncommon with growing season burns in scrub oak communities (Fig. 65). Depending 

on safety and containment issues unique for each individual scrub oak burn, dormant season 

fires may be the wiser choice to lessen the fire intensity problem. The tradeoff is less control 

of encroaching hardwoods. Appropriate fuel models for planning prescribed fires in scrub oak 

communities are AFM-05 or AFM-10 for dormant season burns and AFM-06 or AFM-07 for 

growing season burns. A custom fuel model may also be appropriate because of the presence 

of mountain laurel which is not well represented by any of the standard 13 fuel models. See 

Waldrop and Goodrick (2012) for more details on planning and conducting prescribed fires.

Figure 64.—In the extended absence of fire, scrub oak communities can 

develop too much of a pine component or be invaded by hardwoods such as 

red maple. Photo by Patrick Brose, U.S. Forest Service.
73

EPILOGUE

The role of fire in the upland oak ecosystems of eastern North America is complex. Humans 

have been starting fires for a myriad of reasons throughout eastern North America since the 

end of the last ice age. Those fires have helped establish and perpetuate oak forests, savannas, 

shrublands, and woodlands. Presently, fire is ecologically extinct as an important process, and 

the upland oak ecosystems are in decline because of this absence of fire. Prescribed fire, when 

applied correctly along with other forest management practices, can reverse this decline and 

perpetuate upland oak ecosystems into the future.

ACKNOWLEDGMENTS

The authors are indebted to a multitude of fellow scientists and practicing foresters for 

numerous conversations over the past few years regarding the fire-oak hypothesis and their 

insights into its complexities and intricacies. We especially appreciate their encouragement to 

complete a project that became far more complicated and protracted than we ever imagined. 

Special thanks are due to our reviewers–Margit Bucher, Stacy Clark, Tara Keyser, Greg 

Nowacki, Susan Stout, and two anonymous individuals. We appreciate their time and effort to 

carefully read earlier drafts of this General Technical Report (dubbed “The Behemoth” due to 

its size) and make detailed comments that helped with clarity and conciseness. Additionally, a 

special thank you is due Amy Scherzer who tackled the onerous task of providing the technical 

editing. Finally, we thank the Joint Fire Science Program for funding this project. Without 

the funding and the input of these individuals, this General Technical Report would still be a 

concept, not reality.

Figure 65.—Intense fires are a critical part of the ecology of scrub oak 

communities. Photo by Jennifer Case, The Nature Conservancy.
74

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APPENDIX 1: COMMON AND SCIENTIFIC NAMES OF THE FLORA AND 

FAUNA MENTIONED IN THIS REPORT

Common Name Scientific Name

Flora

American beech Fagus grandifolia

American chestnut Castanea dentata

bear oak Quercus ilicifolia

blackberry Rubus spp.

black cherry Prunus serotina

black oak Quercus velutina

blueberry Vaccinium spp.

bur oak Quercus macrocarpa

cherrybark oak Quercus pagoda

chestnut oak Quercus prinus

common serviceberry Amelanchier arborea

cucumbertree Magnolia acuminate

dwarf live oak Quercus minima

eastern hemlock Tsuga canadensis

eastern white pine Pinus strobus

flowering dogwood Cornus florida

garlic mustard Alliaria petiolata

hayscented fern Dennstaedtia 

 punctilobula

hickory Carya spp.

huckleberry Gaylussacia spp.

loblolly pine Pinus taeda

longleaf pine Pinus palustris

maple Acer spp.

Mohr oak Quercus mohriana

mountain laurel Kalmia latifolia

northern red oak Quercus rubra

oak Quercus spp.

pignut hickory Carya glabra

pin cherry Prunus pensylvanica

pitch pine Pinus rigida

post oak Quercus stellata

red maple Acer rubrum

red pine Pinus resinosa

Common Name Scientific Name

runner oak Quercus pumila

sassafras Sassafras albidum

scarlet oak Quercus coccinea

shortleaf pine Pinus echinata

silver maple Acer saccharinum

striped maple Acer pensylvanicum

sugar maple Acer saccharum

sweet birch Betula lenta

sweet fern Comptonia peregrina

Table Mountain pine Pinus pungens

tree-of-heaven Ailanthus altissima

white oak Quercus alba

witch-hazel Hamamelis virginiana

yellow-poplar Liriodendron tulipifera

Fauna

acorn weevil Curculio spp., 

 Conotrachelus spp.

black bear Ursus americanus

blue jay Cyanocitta cristata

cerulean warbler Setophaga cerulea

eastern chipmunk Tamias striatus

gall wasp Callirhytis spp.

gypsy moth Lymantria dispar

orangestriped oakworm Anisota senatoria

red-headed woodpecker Melanerpes 

 erythrocephalus

red-eyed vireo Vireo olivaceus

ruffed grouse Bonasa umbellus

scarlet tanager Piranga olivacea

tree squirrel Sciurus spp.

Valentine moth Valentina glandulella

white-footed deer mouse Peromyscus leucopus

white-tailed deer Odocoileus virginianus

wild turkey Meleagris gallopavo

wood duck Aix sponsa
Printed on Recycled Paper

Brose, Patrick H.; Dey, Daniel C.; Waldrop, Thomas A. 2014. The fire–oak literature 

of eastern North America: synthesis and guidelines. Gen. Tech. Rep. NRS135. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, 

Northern Research Station. 98 p.

Guidelines for using prescribed fire to regenerate and restore upland oak forests, 

woodlands, and savannas in eastern North America were developed by synthesizing 

the results of more than 100 scientific publications. The first four chapters provide 

background information on the values of oak ecosystems, eastern fire history, oak’s 

adaptations to fire, and the findings of fire-oak research conducted over the past 50 

years. The final chapter synthesizes that background information into guidelines that 

explain how to use prescribed fire to facilitate oak seedling establishment, release 

oak reproduction from competing mesophytic hardwoods, and rehabilitate open oak 

woodlands, oak savannas, and scrub oak communities. A reference section is also 

provided for readers desiring to delve more deeply into the associations between 

periodic fire and oak forests, woodlands, and savannas.

KEY WORDS: fire history, fire-oak hypothesis, prescribed fire, Quercus, sustainable 

forestry

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