Restoration in the 

Southern Appalachians:

A Dialogue among Scientists, Planners, 

and Land Managers

W.T. Rankin and Nancy Herbert, Editors

Forest Service

Research & Development

Southern Research Station

General Technical Report SRS-189

United States Department of Agriculture

2005 2010
March 2014

Southern Research Station

200 W.T. Weaver Blvd.

Asheville, NC 28804

www.srs.fs.usda.gov

The Editors:

W.T. Rankin, Threatened and Endangered Species 

Program Manager, U.S. Department of Agriculture Forest 

Service, Southern Region, 1720 Peachtree Rd. NW, Suite 

700, Atlanta, GA 30309; and Nancy Herbert, Assistant 

Director for Research (retired), U.S. Department of 

Agriculture Forest Service, Southern Research Station, 

200 W.T. Weaver Blvd., Asheville, NC 28804

Cover:

Before-and-after photographs of a restoration project 

on the Buck Creek Serpentine Barrens in western North 

Carolina. Left: Spring 2005 (photo by Paul Davison, 

University of North Alabama). Right: Fall 2010 (photo 

by W. T. Rankin, USDA Forest Sevice)
Restoration in the Southern Appalachians: 

A Dialogue among Scientists, Planners, 

and Land Managers

W.T. Rankin and Nancy Herbert, Editors

iii

CONTENTS

Prologue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iv

1. The Role of Fire in the Southern Appalachians . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Did fire occur in the Southern Appalachians historically? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1

What are the different kinds of fire? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

What are the effects of fire on nongame species in the Southern Appalachians? . . . . . . . . . . . . . . . . . . . . .3

What are the effects of fire on soils in the Southern Appalachians? . . . . . . . . . . . . . . . . . . . . . . . . . . . .4

What are the effects of fire on air quality in the Southern Appalachians? . . . . . . . . . . . . . . . . . . . . . . . . 5

Are there ecosystems in the Southern Appalachians where fire isn’t appropriate? . . . . . . . . . . . . . . . . . . . .7

If we don’t use fire as a management tool, what else do we use? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

How do we restore fire to ecosystems where it has been excluded for many years? . . . . . . . . . . . . . . . . . . 10

What are the consequences if we don’t use fire? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

2. Early Successional Habitat in the Southern Appalachians . . . . . . . . . . . . . . . . . . . . . . . . . . 14

Why are early successional forests important habitats for wildlife species? . . . . . . . . . . . . . . . . . . . . . . 14

What species are favored by early successional habitat and how strong is the relationship? . . . . . . . . . . . . . . 17

Are some wildlife species negatively affected by creating early successional habitat? . . . . . . . . . . . . . . . . . 18

Do tree species differ in their response to early successional habitat? . . . . . . . . . . . . . . . . . . . . . . . . . .18

How much early successional habitat is necessary to provide for biodiversity? 

 What is the optimal percentage of early successional habitat to meet desired goals? . . . . . . . . . . . . . . . 19

What is the natural distribution of early successional habitat across the landscape? . . . . . . . . . . . . . . . . . . 20

Is there a threshold value for early successional habitat patch size? . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

How do we sustain early successional habitat in the landscape? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

Are natural disturbances such as canopy gaps sufficient to maintain early successional habitat? . . . . . . . . . . . 23

Are openings created by fire sufficient to produce early successional habitat? . . . . . . . . . . . . . . . . . . . . . 24

What are the consequences if we do not manage for early successional habitat? . . . . . . . . . . . . . . . . . . . . 24

3. Oak Regeneration in the Southern Appalachians . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

What are the current management practices for regenerating oak forests? . . . . . . . . . . . . . . . . . . . . . . . 25

In the absence of management, can oaks trees regenerate naturally? . . . . . . . . . . . . . . . . . . . . . . . . . . 26

Are there treatments that could improve forest health in oak communities? . . . . . . . . . . . . . . . . . . . . . . 27

What is the role of fire in regenerating oak trees? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

What are the effects of cutting oak trees on high elevation sites? . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

What are the consequences if we do not plan for oak regeneration? . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

Contributing Scientists . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

Indices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

Additional Fire References for the Southern Appalachians . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
iv

PROLOGUE

On April 28, 2009, leaders from the Southern Region (R8) and the Southern Research Station (SRS) of 

the Forest Service, U.S. Department of Agriculture (USDA) met in Asheville, North Carolina, to assess 

the status of restoration efforts in the Southern Appalachian Mountain region. Forest Supervisors from 

the national forests in Virginia, North Carolina, South Carolina, Georgia, Tennessee, and Kentucky 

described restoration efforts underway on their national forests, with a focus on the portion of their 

national forests in the Southern Appalachians. Interspersed in their summaries were comments about the 

need for more research to support a particular restoration technique or restoration goal.

Although many of the topics they discussed have been the subject of considerable research, the 

information was not always in a useful format for Forest Supervisors and their staffs. Former SRS 

Director Jim Reaves offered to summarize the pertinent information into a format that would serve 

their needs. Responding to this request, the group met for a brainstorming session to identify the most 

pressing questions confronting ecosystem restoration in the Southern Appalachians. This session 

produced three areas of focus:

l The role of fire in the Southern Appalachians

l Early successional habitat in the Southern Appalachians

l Oak regeneration in the Southern Appalachians

SRS then worked with R8 Forest Supervisors to develop a list of questions in each of these focus areas. 

A survey of the planning staffs on the six national forests led to a list of additional questions. These 

questions were presented to SRS experts in the three areas of focus. Experts providing responses 

included employees from the Southern Research Station, Northern Research Station, District, Forest 

Health Protection, and Air Resources staffs in the Southern Region, and the University of Tennessee.

This publication uses a question-and-answer format. The questions were posed by Forest Supervisors 

and their planning staffs, and address issues in the restoration of unhealthy or degraded forest 

ecosystems. The answers were provided by conservation scientists from multiple organizations, all of 

whom shared a deep concern about the issues confronting the region. Common and scientific names for 

all species mentioned are listed in the taxonomic index beginning on p. 44.
THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

Did fire occur in the Southern Appalachians 1.1 historically?

Fire has apparently played an integral role in determining 

historical patterns of forest vegetation across the Southern 

Appalachian Mountain region. Fossil pollen and charcoalparticle analyses suggest recurrent fire was common 

in forests of the region for at least 3,000 years before 

the arrival of Europeans (Delcourt and Delcourt 1997, 

Fesenmyer and Christensen 2010). Historical accounts 

suggest anthropogenic fire, often used to affect forest 

structure and composition, was common both before and 

after European colonization (DeVivo 1991, Fowler and 

Konopik 2007, Stewart 2002, Van Lear and Waldrop 

1989). In addition, many of the traits characteristic of plant 

species in the Southern Appalachians can be interpreted as 

evolutionary responses to fire (Christensen 1977, Landers 

1991, Lorimer 1985).

The Five Periods of Anthropogenic Fire Regimes 

Fowler and Konopik (2007) outlined five periods of 

anthropogenic fire regimes in the Southern Appalachians, 

based on changing cultures, population sizes, and land use 

priorities:

Circa 12,000 BP to 1500 AD—During the first period, 

approximately 12,000 BP to 1500 AD, Native Americans 

most likely burned valleys near settlements to clear land for 

agriculture, while upper slopes and ridges were selectively 

burned to promote wildlife habitat. Based on estimates of 

population size and the amount of cleared land necessary 

to support these populations, the spatial effects of Native 

American burning may have reached one-quarter to onehalf the amount of the current farmland in the Eastern 

States (Stanturf and others 2002). Fire return intervals 

varied between 1 and 12 years, depending on elevation, 

slope, aspect, and proximity to native villages (Barden 

1997, Delcourt and Delcourt 1997, Frost 1995).

Circa 1500 to mid-late 1800s AD—The second period of 

fire use began with the arrival of European colonists in the 

16th century. As the number of colonists increased, much of 

the landscape was occupied by settlers who adopted Native 

American practices. Recent dendrochronologies addressing 

this period have documented fire return intervals in xeric, 

central Appalachian oak and pine forests between 5 and 20 

years (e.g., Aldrich and others 2010).

Mid-late 1800s to early 1900s AD—The third period 

of fire coincided with industrialization, beginning in the 

latter half of the 19th century, as railroads improved both 

the access to the mountains and the movement of large 

amounts of commodities. Large-scale timber harvests 

between 1880 and 1920 resulted in heavy fuel loads from 

slash, and created drier, more open stands. Fires were used 

to reduce slash and enhance grazing. The high fuel levels 

from the slash produced much higher intensity fires than in 

previous eras, although the frequency of the fires remained 

similar to earlier periods (Harmon 1982).

1. THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

Restoration in the Southern Appalachians: A Dialogue 

among Scientists, Planners, and Land Managers

W.T. Rankin and Nancy Herbert, Editors

ABSTRACT

We address three key questions for restoration ecology in the Southern Appalachian Mountains. First, what is the role of fire, especially 

when used as a management tool for oak-dominated ecosystems? Second, what is the relationship between early successional habitat 

and biodiversity? And third, how do we regenerate oak ecosystems? To answer these questions, first, we examine the historic role 

of fire in the mountains, discuss its effects on forest resources, and summarize a strategy for restoring fire to ecosystems with a 

long history of fire exclusion. Second, we examine the relationship between early successional habitats and wildlife resources in the 

mountains, discuss the pattern and rate of natural disturbance, and provide suggestions for creating and maintaining early successional 

habitat. And third, we review current management for oak regeneration and discuss the implications for oak ecosystems in the absence 

of management. In addition to addressing current questions in restoration ecology, we provide an extensive bibliography of the 

scientific literature, especially for fire management. Our goal is to provide a concise and practical summary of the current restoration 

literature for use by forest planners and managers throughout the Southern Appalachian Mountains.

Keywords: Early successional habitat, natural disturbance patterns, natural fire, oak regeneration, prescribed fire, restoration ecology, 

Southern Appalachian Mountains, wildlife.
2

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

Early 1900s to late 1900s AD—The fourth period 

of fire began in the early 20th century. Following the 

high-intensity fires of the third period, forest managers 

actively suppressed wildland fire and discontinued 

the use of anthropogenic fire. Fire exclusion, however, 

caused important changes in the structure and function 

of Southern Appalachian forests, especially increases in 

fire-intolerant species, and concomitant decreases in firetolerant species (Vose 2000, 2003).

Late 1900s to present—The fifth period of fire began in 

the late 20th century. A half century of fire suppression 

created forests with heavy fuel loads, creating the potential 

for devastating wildland fires. Beginning in the 1970s, 

forest managers in the Southern Appalachians began using 

prescribed fire, especially in xeric forests dominated by 

pines and oaks, to reduce fuel loads and improve forest 

health. Prescribed fires are now the most common form of 

anthropogenic fire in the Southern Appalachian Mountains.

In summary, fire is a long-standing feature of the Southern 

Appalachian landscape. Although cultural perceptions and 

management practices have changed, especially during 

the past 50 years, all available evidence, from pollen cores 

to dendrochronologies to written and verbal histories, 

suggest fire, either natural or anthropogenic, has played 

an important role in the Southern Appalachians for many 

centuries (Spetich and others 2011).

— Tom Waldrop, Center for Forest Disturbance Science, 

Clemson, SC

— Jennifer Knoepp, Coweeta Hydrologic Laboratory, 

Otto, NC

1.2 What are the different kinds of fire?

Fire can be classified in multiple ways. One of the 

more common classifications is to distinguish between 

prescribed fires and wildfires. Fires can also be classified 

by intensity and season. 

Classifying Fires

Prescribed fire, also known as controlled burning, is fire 

applied to ecosystems, at specific locations, and under 

specific weather conditions, to accomplish predetermined 

management objectives. Fire prescriptions typically control 

effects on ecosystems by controlling fire intensity, either 

by choosing the proper environmental conditions—wind, 

humidity, fuel moisture—or through site preparation. Fire 

prescriptions also address fire behavior and spread, by 

specifying the movement of the fire relative to the wind: 

head fires spread in the direction of the wind, backing 

fires spread against the wind, and flanking fires spread at 

right angles to the wind. Because wind patterns and fuel 

conditions are more variable in the mountains than in other 

regions of the South, considerable experience and training 

are required to conduct a successful prescribed fire in the 

Southern Appalachians (e.g., Achtemeier 2008).

Wildland fires, on the other hand, are unplanned. Although 

prescribed fires and wildland fires can share many 

characteristics, wildland fires are more likely to burn under 

severe fuel and weather conditions, creating hot fires that 

are difficult, and dangerous, to control. Because they are 

more likely to burn hot, wildland fires are also more likely 

to adversely affect Southern Appalachian forests, killing 

desirable trees and consuming the organic portion of the soil.

Some wildland fires are used to achieve management 

objectives. These fires, although not started by humans, are 

allowed to burn under specific fuel and weather conditions. 

In the Southern Appalachians, these fires are generally 

limited to wilderness areas or lands managed by the 

National Park Service.

Low-intensity fires rarely have the same effects as highintensity fires. For example, less intense fires are less likely 

to produce early successional habitat than hotter, more 

intense fires. The effects of fire intensity, however, also 

depend upon the season. In general, dormant-season fires in 

the Southern Appalachians are more intense than growingseason fires, because growing-season moisture, combined 

with high humidity, often suppresses fire intensity. The 

effects of low-intensity fires during the growing season, 

however, can be similar to, or even more severe than, the 

effects of high-intensity fire during the dormant season, 

because the stem of most woody plants is severely damaged 

when the cambium layer reaches 145 °F (Wright and Bailey 

1982), and this temperature is more easily reached during 

the heat of the growing season. In addition, most of the 

carbohydrates in shrubs and trees are located aboveground 

(Knapp and others 2009), so growing-season fire typically 

kills woody species more effectively than dormant-season 

fires. When these plants are topkilled, the plant contains 

fewer reserves for resprouting (Drewa and others 2002).

Early results from ongoing research suggest that multiple 

growing-season burns reduce woody cover while increasing 

herbaceous cover.1

 In general, however, the effect of 

growing-season fire on plant and animal communities in the 

Southern Appalachians is poorly documented, and not well 

understood (see Knapp and others 2009). This is especially 

true for long-term effects over a range of fire intensities 

(Fontaine and Kennedy 2012).

1 Unpublished data. On file with: Craig Harper, Professor of Wildlife 

Management and the Extension Wildlife Specialist at the University of 

Tennessee, 280 Ellington Plant Sciences Building, Knoxville, TN 37996.
3

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

Fire Effects 

Prescribed fires are used for a wide variety of objectives 

throughout the South. They can reduce hazardous fuels, 

dispose of logging debris, prepare sites for seeding or 

planting, influence vegetation composition and structure 

for many wildlife species, manage competing vegetation, 

control insects and disease, and improve forage for grazing. 

Scientists at Coweeta Hydrologic Laboratory have studied 

three forms of prescribed fire treatments: fell and burn, 

stand replacement, and understory. Each of these approaches 

has different ecological effects, and choosing among them 

depends on the desired future condition for the ecosystem.

Fell-and-burn—In the fell-and-burn approach, the 

ecosystem is thinned, and the slash cured in place for 

several months. This process increases the amount of 

coarse fuel in the ecosystem, which, in turn, increases 

the fire intensity. The fell-and-burn technique has been 

applied to pine-hardwood ecosystems, where it increases 

the productivity and insect resistance of commercial tree 

species (Vose 2000).

Stand-replacement—Stand-replacement fires are ignited 

during periods of low fuel moisture and high winds. These 

conditions increase the intensity of the fire to a point where 

the overstory trees are killed, allowing the establishment 

of a new, or replacement, stand. Stand-replacement fires 

produce a mosaic of fire effects across the landscape, due to 

spatial heterogeneity in fire intensity. Compared to the felland-burn technique, stand-replacement fires are less likely 

to impact the biogeochemical cycle—for example, nitrogen 

in the soil, carbon on the forest floor, and the chemistry of 

the streams—with most of the losses occurring on ridges 

(Vose and others 1999).

Understory—Understory fires are ignited during periods 

of high fuel moisture and low winds, limiting the effects 

to the fine fuels, such as small twigs and fallen leaves, 

on the forest floor. Understory fires are used to reduce 

fuels and influence the composition and structure of 

understory vegetation, which indirectly affects food and 

cover resources for many wildlife species (Jackson and 

others 2007, McCord and Harper 2011). Several researchers 

have shown understory fires in xeric, intermediate, and 

mesic sites increase the diversity of understory plants, with 

no measurable negative impacts on water quality or site 

productivity (Elliott and Vose 2005b, Elliott and others 

2004, Hubbard and others 2004, Vose 2003).

— Tom Waldrop, Center for Forest Disturbance Science, 

Clemson, SC

— Jennifer Knoepp, Coweeta Hydrologic Laboratory, 

Otto, NC

— Craig Harper, University of Tennessee, Knoxville, TN

What are the effects of fire on nongame 1.3 species in the Southern Appalachians?

Fire effects on wildlife are most closely associated with 

changes to habitats and microhabitats in the forest, such as 

changes to the trees, shrubs, and leaf litter. Low-intensity 

burns generally do not kill trees. Because the trees are 

not killed, the general structure of the forest remains 

unchanged, and microhabitats within the stand either are 

little affected or recover quickly. In contrast, hot fires that 

kill trees can change the forest structure, significantly 

altering light levels and microclimates throughout the 

stand, and therefore may have a substantial effect on 

wildlife communities. In the Southern Appalachians, most 

prescribed fire is low intensity and does not significantly 

alter the general structure of the stand.

Small mammals—The few published studies examining 

the effect of burns on small mammals are inconsistent, 

likely due, in part, to high variability among the sites, and 

low levels of replication within the studies. Some studies 

report similar white-footed or deer mouse (Peromyscus

spp.) abundance in both burned and unburned hardwood 

forest (Ford and others 1999, Keyser and others 2001, 

Raybuck and others 2012), while others (Kirkland and 

others 1996) report a lower abundance of white-footed 

mice in burned stands. In contrast, Krefting and Ahlgren 

(1974) reported higher densities of deer mice in burned, 

mixed conifer-hardwood forest sites. In the Southern 

Appalachians, we found more white-footed mice in burned 

stands, possibly because the fire reduced the depth of 

the leaf litter (Greenberg and others 2006). Increased 

populations of Peromyscus on burned sites have been 

attributed to better visibility and higher concentrations of 

seed, a food source for the mice, after reductions in litter 

cover and depth (Ahlgren 1966, Tester 1965).

Bats—Prescribed fire may have direct, short-term effects 

on bats as well as indirect, long-term effects. Short-term 

effects include disturbance, and potential mortality, of 

litter-hibernating bats such as eastern red bats during 

winter burns (Moorman and others 1999, Saugey and 

others 1989), and disturbance of crevice-roosting bats such 

as northern long-eared and Indiana bats during late spring 

and summer burns (Dickinson and others 2009). Fire 

may also destroy roosting sites, temporarily reducing the 

suitability of burn sites for roosting (Moorman and others 

1999, O’Keefe and Loeb 2010).

Most studies in the Eastern United States, however, have 

found the long-term effects of prescribed fire are either 

neutral or beneficial to bats. Because fire often reduces 

physical obstructions in the environment that may 

interfere with echolocation and maneuver, bat foraging 

activity often increases in burned areas (Lacki and 
4

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

others 2009, Loeb and Waldrop 2008, Smith and Gehrt 

2010). Insect abundance may also increase following fire, 

producing more suitable foraging sites (Lacki and others 

2009, Malison and Baxter 2010).

Prescribed fire may also improve summer roosting habitat. 

Northern long-eared bats (Lacki and others 2009) and 

evening bats (Boyles and Aubrey 2006) select roost trees 

in burned sites, perhaps due to increased solar radiation. In 

addition, male Indiana bats appear to select roosts in both 

pine trees (MacGregor and others 1999) and hardwood 

trees (Johnson and others 2010) in burned sites. The effects 

of fire on Indiana bat maternity colonies in the Southern 

Appalachians are currently being investigated (O’Keefe 

and Loeb 2010).

Herpetofauna—Among the few studies of the 

herpetofauna in the Southern Appalachians, most suggest 

prescribed fire does not substantially change amphibian 

abundance. For example, Ford and others (1999) reported 

high-intensity prescribed fire had no effect on woodland 

salamanders. Matthews and others (2010) found lowintensity prescribed burns that did not kill overstory trees 

had no effect on salamanders, and we found similar results 

in a study at the Cold Mountain Game Lands (North 

Carolina), where a prescribed burn had no detectable 

results on terrestrial salamanders (Raybuck and others 

2012). In contrast, we have also found that although a 

single prescribed burn that killed overstory trees did not 

reduce the relative abundance of terrestrial salamanders 

(Greenberg and Waldrop 2008), salamander abundance 

declined after a second burn in the same study sites 

(Matthews and others 2010).

Several studies indicate prescribed fire does not affect the 

abundance of frogs, and may increase toad abundance 

(Greenberg and Waldrop 2008, Matthews and others 

2009). Hot fires that produce overstory mortality may 

benefit reptiles, particularly lizards, by creating more 

open conditions and warmer temperatures (Greenberg 

and Waldrop 2008, Matthews and others 2010, Moorman 

and others 2011, Moseley and others 2003, Renken 2006, 

Russell and others 1999).

Birds—In the Southern Appalachians, we found lowintensity, dormant-season burns had few detectable effects 

on breeding birds (Greenberg and others 2007b). In 

contrast, total bird species richness and density increased 

in communities treated with hot prescribed fire that killed 

trees and created snags. Responses to prescribed fire 

differed considerably among bird species, according to 

their associations with specific habitat features. Because 

the fire removes the leaf litter and shrub layers, bird species 

associated with these structural features may show shortterm decreases in abundance. For example, ground-nesting 

worm-eating warblers, or shrub-dwelling hooded warblers, 

may decline until the leaf litter is replenished the following 

autumn, or until shrub cover increases within a year or two. 

In general, however, high-intensity burning with heavy 

tree-kill can be used to increase bird species associated 

with open habitats, such as indigo buntings and eastern 

bluebirds, or snags (woodpeckers and secondary-cavity 

nesters), while retaining many forest and generalist species 

(Greenberg and others 2007b).

— Katie Greenberg, Bent Creek Experimental Forest, 

Asheville, NC

— Susan Loeb, Clemson University, Clemson, SC

What are the effects of fire on soils in the 1.4 Southern Appalachians?

Due to slow rates of decomposition, brought on by 

a combination of dry conditions and low nutrient 

concentrations, fire-dependent forests tend to accumulate 

woody debris and plant materials on the forest floor (e.g., 

Woodall and Likens 2008). As a result, fire can play an 

important role in these stands, quickly returning nutrients 

to the mineral soil, and creating a more open environment 

that allows seed germination and seedling establishment.

Much of the research on fire effects to forest soils has 

focused on soil organic matter. Soil organic matter is 

critical for maintaining soil quality because it retains water, 

carbon, and essential nutrients, such as nitrogen. 

A meta-analysis of more than 40 published papers showed 

that, on average, prescribed burning had no effect on total 

soil carbon and nitrogen concentrations (Johnson and 

Curtis 2001). The effect of fire on soil carbon and nutrients, 

however, varied greatly with site and burning conditions. 

In general, higher intensity burns reduce soil organic 

matter by directly consuming organic carbon, as well as 

soil nutrients, by volatilizing organic nitrogen (Knoepp 

and others 2005). High-intensity burns also consume 

large amounts of the organic materials on the forest floor, 

removing future soil organic matter, or exposing mineral 

soil, potentially increasing both soil erosion and nutrient 

loss (Robichaud and Waldrop 1994). On the other hand, 

burning woody debris and plant material on the forest 

floor can add cations such as calcium and magnesium to 

the surface soil, increase soil pH, and increase nitrogen 

availability, especially if some of the volatilized nitrogen 

is retained (Kovacic and others 1986, Raison and others 

1990). In addition, low-intensity burns appear to have 

little effect on soil erosion, even in the steep terrain of 

the Southern Appalachians, due to rapid regrowth of the 

burned community (Van Lear and Danielovich 1988).
5

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

Most research on fire effects in the Southern 

Appalachians has examined the impacts of single fires. 

In comparison, the effects of multiple fires over a period 

of years are poorly studied. Although most soil variables 

show little change immediately after burning (Knoepp 

and others 2004), recurring fire may produce cumulative 

effects not evident after a single fire. For example, Vance 

and Henderson (1984), working in Missouri on an oak 

flatwoods burned repeatedly over 30 years, found nitrogen 

mineralization was reduced by long-term burning, and 

Neary and others (2003), working in the Western United 

States, found that shorter fire return intervals may reduce 

carbon and nitrogen in surface soils. Long-term burning 

may also result in an accumulation of recalcitrant forms 

of carbon, including black carbon (Ponomarenko and 

Anderson 2001).

Only one study, the National Fire and Fire Surrogate Study 

(e.g., Youngblood and others 2007), has shown impacts to 

soils after repeated burning in the Appalachian region. The 

study encompassed two mid-elevation, oak-hickory forest 

communities in the Appalachian plateau of southeastern 

Ohio, with two dormant-season fires on each site. Principal 

effects included the following:

l Direct soil heating, which may alter soil properties and 

kill soil organisms (Boerner and others 2005, Giai and 

Boerner 2007), but did not increase soil compaction 

(Boerner and others 2007);

l The volatilization and convection, in ash, of nitrogen, 

phosphorus, and cations (Coates and others 2008, 

Huang and others 2007);

l Minor levels of mineral soil exposure, which may lead 

to sheet erosion if the slope is sufficient (Coates 2006);

l Subtle and transient changes in pH, nitrogen availability, 

organic carbon, C/N ratio, and soil microorganisms 

(Boerner and others 2006, 2007; Coates and others 2008);

l Complex effects due to the interactions of site quality, 

slope position, and fire behavior (Boerner 2006).

Boerner and others (2007) concluded that both prescribed 

fire and restoration thinning can be applied to oak-hickory 

forests without significant negative effects on forest soils.

In summary, high-intensity fires, such as the fires used to 

prepare sites for planting, may reduce soil organic matter, 

reducing the amount of soil nutrients, and increase the 

amount of exposed mineral soil, increasing the risk of 

erosion. When properly applied, however, low-intensity, 

prescribed fire minimizes these effects, producing 

little change in soil nutrients (Boerner and others 2007, 

Knoepp and others 2004) and soil erosion (Van Lear and 

Danielovich 1988).

— Tom Waldrop, Center for Forest Disturbance Science, 

Clemson, SC

— Jennifer Knoepp, Coweeta Hydrologic Laboratory, 

Otto, NC

What are the effects of fire on air quality in 1.5 the Southern Appalachians?

The primary products of burning forest fuels are heat, 

water, and carbon dioxide. Wildland fires, however, rarely 

burn fuels completely. As the temperature of the fire drops, 

fuels begin to smolder, producing smoke. Depending on 

the amount and condition of the fuels, all woodland fires 

produce smoke to some degree. Smoke is a complex mixture 

of toxic gases, including carbon monoxide, sulfur dioxide, 

nitrogen dioxide, and ozone, as well as particulate matter, 

including soot and tar (Chi and others 1979). The effects of 

wildland fire, therefore, can be related to either the direct 

effects of the toxic gases on human health, or the indirect 

effects of the particulates on the human environment, 

especially in relation to fog and traffic safety. When any 

component of smoke is present at levels that adversely affect 

human health and safety, it can be considered a pollutant.

In general, wildland fire, especially prescribed fire, rarely 

produces gases at levels affecting human health. Less 

than 3 percent of the total national emissions of carbon 

monoxide and hydrocarbons can be attributed to prescribed 

fire (SAMAB 1996). Wildland fire is not likely to release 

nitrogen oxides in significant quantities because the 

threshold temperature for the release of these compounds, 

2,700 °F, is hotter than the temperatures that normally 

occur during prescribed fires (McMahon and Ryan 1976). 

Compared to natural and anthropogenic sources, sulfur 

dioxide emissions from forest fires are negligible (Hall 

1972). Most atmospheric sulfur dioxide comes from natural 

sources, such as volcanoes, oceans, and plant decay; 

industrial sources account for about 10 percent of total 

emissions (Hardy and others 2001, Komarek 1970).

Small amounts of nitrogen oxides and volatile organic 

compounds, however, are precursors to ground-level 

ozone and may become important in locations where 

ozone levels are already problematic. These locations 

include low-elevation (<3,500 feet), intermountain basins 

in the Southern Appalachians, especially during the 

summer months, when high-pressure weather systems 

limit the amount of upper-level mixing. In addition, 

sunlight working directly on nitrous oxide compounds 
6

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

can form other compounds, primarily ozone, that are 

potentially harmful to vegetation, particularly during 

the growing season. For example, in February 2007, 

two prescribed fires in central Georgia increased 

ozone concentrations in the Atlanta metropolitan area 

approximately 30 percent. These elevated levels were 

primarily due to increases in nitrogen oxides and volatile 

organic compounds released from the fuels, but also due 

to increases in volatile organic compounds released by 

the heating of the forest canopy (Hu and others 2008, Liu 

and others 2009). Most wildland fires in the Southern 

Appalachians, however, occur during the spring and fall, 

when windy weather produces dry fuels, favoring more 

efficient combustion. Windy conditions also improve 

atmospheric mixing, which dilutes emissions over a 

wider area. As a result, ozone tends to rapidly disperse 

to acceptable levels. Because dispersal is effective at 

managing ozone levels, many of the potentially negative 

effects of wildland fires on air quality can be mitigated by 

carefully planning and executing controlled burns.

The major pollutant produced by prescribed burning is 

particulate matter (Dieterich 1971, Sandberg and others 

1979). Particulate matter is a complex mixture of soot, 

tars, and volatile organic compounds (McMahon 1977). 

Although fires produce many sizes of particles, the 

most problematic are the smallest particles, typically 

defined as <2.5 microns in diameter, because they can 

remain airborne for a considerable amount of time, and 

they can penetrate deeply into human lungs where they 

may contribute directly to respiratory problems (Hardy 

and others 2001). Particles <2.5 microns represent 

approximately 70 percent of the particles in smoke from 

wildland fires (Hardy and others 2001).

Concentrations of particulate matter are tracked by the 

Forest Service and compared to the National Ambient Air 

Quality Standard (U.S. Forest Service 2011). Wildland fires 

in western North Carolina have occasionally exceeded the 

standard for particles ≤2.5 microns, primarily due to heavy 

fuel loads produced by outbreaks of southern pine beetle. 

Compared to gaseous and particulate atmospheric pollutants 

from industrial, utility, or mobile sources, however, wildland 

fires are relatively minor (SAMAB 1996). In general, 

the concentration of particles <2.5 microns produced by 

prescribed burning does not exceed the National Ambient 

Air Quality Standard (Hardy and others 2001).

On the other hand, public concern is likely to occur before 

particulate levels violate National Ambient Air Quality 

Standards. For example, particulates also affect visibility. 

Most of the haze in the Southern Appalachians can be 

attributed to particles ≤ 2.5 microns, particularly in the 

morning, when humidity tends to be high (Abdel-Aty 

and others 2011). When wind speed is low and humidity 

high, moisture in the air condenses around particulates, 

forming dense smoke or a combination of smoke and fog. 

This potentially leads to traffic safety issues, and the risk 

of smoke moving into sensitive areas such as airports, 

highways, and communities is probably the major concern 

related to air quality and prescribed burning. Smoke that 

disperses during the night, when relative humidity is near 

100 percent, can also result in superfog, a condition where 

visibility is reduced to a few feet (Achtemeier 2009). On 

a busy highway, superfogs can contribute to multiple 

collisions and traffic pileups. For example, in January 2008, 

a superfog from a woodland fire in central Florida led to a 

70-car pileup that killed 4 motorists on Interstate 4.

Smoke also contributes to regional haze, reducing visibility 

at scenic views (Tombach and Brewer 2005). Recently, the 

Forest Service participated in a technical analysis aimed at 

reducing fire emissions and regional haze in the Southern 

Appalachians (SAMI 2002). Currently, the emissions from 

wildland fires are not considered a significant contributor to 

regional haze (SAMAB 1996).

Problem smoke is a chronic issue in the South for several 

reasons. First, prescribed fire produces large amounts of 

smoke. To avoid damaging tree roots, prescribed burning 

is typically conducted when soil and litter are moist. Moist 

fuels burn less efficiently and smolder longer than dry 

fuels, both increasing the amount of smoke produced and 

reducing the amount of heat available to carry the smoke 

aloft. This combination, in turn, increases the likelihood 

that the smoke will stay close to the ground. Second, the 

climate is characterized by air masses that trap smoke 

close to the ground. During the winter, shallow valleys 

can develop atmospheric inversions overnight, trapping 

smoke near the ground, and drainage winds can carry 

smoke as much as 10 miles, far enough to reach roadways 

in many locations. Finally, the region is densely populated, 

producing a large amount of wildland-urban interface and 

exacerbating interactions between smoke and the human 

environment.

Because of the potentially serious effects of prescribed fire 

on air quality, guidelines for smoke management have been 

developed by the Forest Service to reduce the atmospheric 

impacts of prescribed fire (Hardy and others 2001). 

Although prescribed fire produces the same emissions 

as wildland fire, forest managers can choose the season 

and weather conditions for prescribed fire, allowing them 

to mitigate many of the effects of the fire. For example, 

dormant-season burns occur at lower fuel moistures and 

under more consistent weather conditions than growingseason burns, producing lower emissions and improving 

control over smoke movements.
7

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

In addition, the Forest Service uses several computer 

models to predict the movement of smoke across the 

southern landscape. For example, VSMOKE models the 

daytime movement and concentration of particulate matter 

in smoke, assuming level terrain and unchanging winds 

(Lavdas 1996). A second application, the PB-Piedmont 

model, models the movement of smoke trapped near 

the ground at night across the complex terrain of the 

Piedmont (Achtemeier 2001). Researchers are planning 

two sister programs of the PB-Piedmont model, one for the 

Appalachians, and one for coastal areas influenced by sea/

land circulations.

Most Southern States require a permit for prescribed 

burning. The permitting process provides a method for 

managing smoke in critical areas, and a mechanism 

for administering laws and other regulations (Brenner 

and Wade 1992, Haines and Cleaves 1995, Yoder and 

others 2003). The permit process is particularly useful 

for maintaining regional air quality standards because it 

allows State agencies to coordinate and evaluate the total 

amount of smoke produced on a regional basis. Obtaining 

a permit may be difficult in some areas of the Southern 

Appalachians because favorable days for controlled 

burning may be few, and many resource managers may 

want to burn when favorable conditions occur. In general, 

however, well-planned controlled burns performed 

under favorable weather and with good estimates of fuel 

consumption and fire intensity, can be implemented 

with a high confidence that air quality standards will be 

maintained (Hardy and others 2001).

— Tom Waldrop, Center for Forest Disturbance Science, 

Clemson, SC

— Bill Jackson, National Forests of North Carolina, 

Asheville, NC

— W. Henry McNab, Bent Creek Experimental Forest, 

Asheville, NC

— Roger Ottmar, Pacific Northwest Research Station, 

Seattle, WA

Are there ecosystems in the Southern 1.6 Appalachians where fire isn’t appropriate?

The appropriate use of fire depends on a variety of factors, 

each dependent on the ecosystem in question:

l What is the history of fire in the ecosystem? For 

example, dry or xeric communities in the Southern 

Appalachians historically exhibited frequent, but lower 

intensity, fires. More mesic forests, on the other hand, 

historically exhibited less frequent, but potentially higher 

intensity, fires (Stanturf and others 2002: Table 25.1).

l What is the fuel condition of the ecosystem? For 

example, dry, shrubby forests have higher fuel loads 

than more mesic, open forests, and are more likely to 

have carried fire in the past. Fuel loads can also change 

in response to fire suppression, overstory disturbance, 

and the presence of invasive plant species.

l What are the perceived effects on the ecosystems? For 

example, fire can substantially change the biodiversity 

and structure of a forest.

The following ecosystems occur in the Southern 

Appalachians, and are therefore potentially subject to 

prescribed fire. For each ecosystem, Reilly and others 

(2012) have assessed the fuel loads and the potential effects 

of prescribed fire.

Bottomland hardwood forests—Bottomland hardwood 

forests are found at the lowest elevations in the major 

river valleys. These forests are very productive, with rapid 

decomposition rates due to seasonal flooding and high soil 

moisture. Floods play an important role in the disturbance 

regime, and may redistribute coarse woody debris and 

remove litter, especially after large events.

Floodplain forests are particularly prone to invasion by 

exotic species (Brown and Peet 2003), and these species 

have potentially altered the fuel structure in bottomland 

hardwood forests. For example, dense thickets of Chinese 

privet and multiflora rose may form large patches of 

continuous fuels capable of carrying fire under dry 

conditions, and kudzu may reach into forest canopies 

along forest edges, creating ladder fuels. The presence 

of invasive species may warrant the use of fire to reduce 

localized fire hazards.

On the other hand, the role of fire in these ecosystems 

is poorly understood. Wade and others (2000) caution 

the tree species associated with bottomland forests tend 

to be sensitive to fire, and the species patterns in these 

communities tend to reflect the hydrology of the system, 

not the fire regime. As a result, bottomland forests in the 

Southern Appalachians do not appear to be fire-adapted 

ecosystems suitable for prescribed burning.

Oak forests—Oak forests are the most extensive 

ecosystems in the Southern Appalachians, occurring across 

a wide range of elevations, and varying in both topography 

and moisture regime. Xeric oak forests are frequently 

dominated by chestnut and scarlet oaks, while mesic oak 

forests are dominated by white oak and northern red oak. 
8

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

A thick layer of potentially flammable shrubs, primarily 

mountain laurel, blueberry, and huckleberry, is often 

present in oak forests, especially in more xeric conditions 

(Waldrop and others 2007). Shrubs can represent a large 

proportion of the hazardous fuels in the community, 

particularly when composed of mountain laurel, and 

frequently pose a serious problem for fuel management 

(Stanturf and others 2002, Waldrop and Brose 1999).

Most studies show only limited benefits to oak following 

prescribed fire (Alexander and others 2008, Hutchinson 

and others 2005b, Signell and others 2005, Wendel and 

Smith 1986). Although the relationship between oak 

regeneration and fire is complex, controlling competing 

vegetation and modifying light in the understory are 

necessary to maintain oak forests in the face of succession 

towards a more mesic condition, in which stands currently 

dominated by oaks would be replaced by stands dominated 

by species such as red maple (Nowacki and Abrams 2008). 

In addition, prescribed fire appears to increase herbaceous 

cover and diversity in the understory of oak forests (Burton 

and others 2011, Elliott and others 2011, Hutchinson 2006, 

Hutchinson and others 2005a).

Because of the historical role of fire in creating and 

maintaining healthy oak forests, prescribed burning can be 

a valuable management tool in these ecosystems.

Southern yellow pine forests—Southern yellow pine 

forests occur on the xeric upper slopes and ridges of the 

Southern Appalachians, at low and middle elevations. 

Dominant species include Virginia pine, pitch pine, 

shortleaf pine, and Table Mountain pine. A dense shrub 

layer, including blueberry, huckleberry, and mountain 

laurel, is frequently present.

Many yellow pine stands were established early in the 

20th century before the period of fire exclusion (Brose and 

Waldrop 2006) and have become decadent (Williams and 

others 1990). Prescribed fire has been commonly used to 

promote regeneration in yellow pines, especially Table 

Mountain and pitch pines, by reducing the number of 

encroaching shrubs and hardwood species. At one time, 

regeneration was associated with intense, stand-replacing 

fires, but more recent research suggests periodic surface 

fires of moderate intensity may be sufficient (Brose and 

Waldrop 2006, Waldrop and Brose 1999).

Southern yellow pine ecosystems represent one of the 

most challenging situations for fuel managers. Flammable 

evergreen canopies and abundant vertical fuels, such 

as mountain laurel, can result in severe crown fires. 

In addition, disturbance such as wind, ice storms, 

and southern pine beetle infestations can increase the 

abundance of both small- and large-diameter woody 

fuels (Waldrop and others 2007). Periodic surface fires 

would not only facilitate regeneration but would also 

reduce dangerous fuel loads. As a result, southern yellow 

pine ecosystems appear ideally suited to a program of 

prescribed fire.

Mixed mesophytic/rich cove forest—Mixed mesophytic 

forests, also known as rich cove forests, are among the most 

diverse communities in the Southern Appalachians. These 

forests are typically found on moist, east- and north-facing 

slopes and sheltered coves, at low and mid-elevations. The 

forests are dominated by yellow poplar, sweet birch, sugar 

maple, and black cherry, and generally support a diverse 

herbaceous flora.

Because they occur in sheltered coves that collect and 

retain moisture, rich coves are generally more mesic than 

other mid-elevation forest communities in the Southern 

Appalachians, with higher fuel moistures. These conditions 

generally reduce the frequency of fire, and rich coves are 

usually associated with low fire frequencies, although few 

studies have examined this relationship closely. Disturbance 

in cove forests is more often associated with canopy 

gaps produced by the fall of one or a few trees (Runkle 

1982, 1990). Periods of prolonged drought can exacerbate 

overstory mortality, which may increase surface fuels and 

midstory density, especially in canopy gaps, increasing the 

possibility of catastrophic fire (Olano and Palmer 2003). As 

a result, prescribed fire may help reduce the likelihood of 

devastating fire in these ecosystems. Compared to oak and 

pine ecosystems, however, the role of fire in rich cove forests 

has been rarely studied, and remains poorly understood 

(Wade and others 2000). In general, rich cove forests do not 

appear suitable for a program of prescribed fire.

White pine-hemlock-hardwood forests—White pinehemlock-hardwood forests are typical of cool, moist 

ravines over a range of elevations. Also known as acidic 

coves, these forests are often composed of large-diameter 

trees at low density, with a thick shrub layer of rosebay 

rhododendron.

The historical disturbance regime of white pine-hemlockhardwood forests was likely dominated by wind (Lorimer 

1995). Although generally long-lived, white pine and eastern 

hemlock are both characterized by shallow root systems, 

and are therefore susceptible to windthrow. These forests 

are also characterized by low fire frequencies because they 

characteristically grow in ravines with high humidity and soil 

moisture. When fire does occur, however, mortality can be 

high (Reilly and others 2006).

The recent invasion of the hemlock woolly adelgid has 

devastated hemlock communities throughout the Southern 

Appalachians. High rates of mortality will likely cause a 
9

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

pulse in both small and large surface fuels (e.g., Waldrop 

and others 2007). Because of these large fuel loads, white 

pine-hemlock-hardwood communities in the Southern 

Appalachians may be unusually susceptible to catastrophic 

fire at the present time.

Given the unusually high fuel loads, the infrequent fire 

return intervals, and the general resistance of the ecosystem 

to burning, white pine-hemlock-hardwood forests do not 

appear suitable for prescribed fire.

Northern hardwood forests—In the Southern 

Appalachians, northern hardwood forests occur in coves, 

and on upper slopes at elevations above 4,000 feet. These 

stands are dominated by hardwood species characteristic 

of northern forests, such as beech, sugar maple, and 

yellow birch. The understory tends to be moist, and 

dominated by ferns.

Disturbance in northern hardwood forests is primarily due 

to wind (Lorimer and Frelich 1994). High rainfall and soil 

moisture keep fuel moisture relatively high, and fire has 

probably been infrequent, with return intervals between 300 

and 500 years (Lorimer 1977). Because of the infrequent 

fire intervals and the overall resistance of the ecosystem to 

burning, fire does not appear to be an important element of 

these forests, and northern hardwood forests do not appear 

suitable for prescribed fire.

Spruce-fir forests—Spruce-fir forests occur at the highest 

elevations in the Southern Appalachians, generally above 

5,000 feet. These forests are dominated by Fraser fir and 

red spruce, with thick litter and relatively few understory 

plants. Growing seasons are short, and the weather is 

characterized by abundant moisture, high humidity, and 

frequent cloud cover. The disturbance regime includes 

wind and ice storms.

These forests are structurally similar to boreal forests, 

and large, high-severity fires may occur during periods of 

prolonged drought (White and others 1985). More recently, 

acid precipitation and balsam woolly adelgid infestations 

have resulted in large-scale mortality of canopy trees, 

creating hazardous fuel conditions, and areas disturbed 

by ice or the adelgid may contain abundant coniferous 

regeneration capable of carrying intense fire (Smith and 

Nicholas 2000). On the other hand, fire frequency is very 

low, with estimated return intervals reaching into the 

millennia (White and others 1985). Given the low fire 

frequency and the extremely moist conditions, spruce-fir 

forests do not appear suitable for prescribed fire.

In general, therefore, fire has played an important role 

in many Southern Appalachian ecosystems, but these 

ecosystems tend to be the more xeric ones at mid-elevations, 

typically dominated by oaks or pines. Because the more 

mesic ecosystems in the Southern Appalachians, such 

as cove forests, are still subject to catastrophic fire under 

drought conditions, prescribed burning may be useful, under 

carefully controlled conditions, to minimize the likelihood 

of catastrophic fires in the future. In general, however, as 

ecosystem moisture increases, and elevations approach 

the extremes of the Southern Appalachian landscape, fire 

becomes increasingly less frequent, more likely to occur 

during drought, more catastrophic when it occurs, and 

therefore increasingly less suitable as a management tool.

— Tom Waldrop, Center for Forest Disturbance Science, 

Clemson, SC

— Tara Keyser, Southern Research Station, Asheville, NC

— Duke Rankin, USDA Forest Service, Southern Region, 

Atlanta, GA

If we don’t use fire as a management tool, 1.7 what else do we use?

Some desirable changes to Southern Appalachian 

ecosystems are better achieved by treatments other 

than fire. For example, bat activity (Loeb and Waldrop 

2008) and diversity (Leput 2004) are more likely to be 

improved by overstory thinning than a single prescribed 

fire. Thinning also increases coarse woody debris on 

the forest floor (Waldrop and others 2004), which may 

improve habitat for small mammals. High-intensity fire 

can accomplish the same objective by killing overstory 

trees (Waldrop and others 2010), but with less predictable 

results.

On the other hand, fire, especially low-intensity, highfrequency fire, has been a component of the Southern 

Appalachians for many years (Wade and others 2000). 

Many of our native ecosystems are fire-adapted, especially 

drier forests dominated by pines or oaks, and these 

forests have proven to be unstable in the 80 years since 

fire suppression became widespread in the Southern 

Appalachians (Nowacki and Abrams 2008). Ultimately, the 

role of fire in restoring and maintaining native ecosystems 

is unique:

l Fire kills and consumes a portion of the aboveground 

vegetation with very little impact to the mineral soil.

l Fire rapidly recycles nutrients back into the ecosystem.

l Fire improves seed germination by removing thick 

layers of duff and coarse fuels.
10

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

l Fire selectively removes fire-intolerant species, restoring 

ecosystem composition and structure.

l Fire improves wildlife value for many game species in 

the Southern Appalachians, including white-tailed deer, 

wild turkey, and ruffed grouse.

l Fire addresses restoration goals by increasing 

biodiversity, including understory plants (Waldrop and 

others 2008), reptiles (Kilpatrick and others 2010), and 

pollinating insects (Campbell and others 2007).

Other stand-level treatments, such as thinning overstory 

trees, felling understory shrubs, and herbicide applications, 

can mimic the changes to ecosystem structure obtained from 

prescribed burning, but all of these treatments add fuels to the 

forest floor, and greatly increase the risk of severe wildland 

fire over a period of ≥5 years (Waldrop and others 2010).

In summary, the combination of selective responses between 

fire-tolerant and intolerant species, the removal of forest 

floor material to allow seed germination, and the addition of 

nutrients to surface soil cannot be fully replicated using any 

other fuel treatment or management action. Returning fire to 

fire-adapted ecosystems is vital to restoring, and improving, 

the health of our native ecosystems.

— Tom Waldrop, Center for Forest Disturbance Science, 

Clemson, SC

— Jennifer Knoepp, Coweeta Hydrologic Laboratory, 

Otto, NC

How do we restore fire to ecosystems where 1.8 it has been excluded for many years?

Fire has a long history in the Southern Appalachian 

landscape. Soil charcoal, tree-ring scars, and fire-adapted 

vegetation all provide evidence for the role of fire as 

a natural process over the past several thousand years 

(Aldrich and others 2010, Fesenmyer and Christensen 2010, 

Flatley and others 2013, Zobel 1969). Beginning in the 

early 20th century, however, land managers in the Southern 

Appalachians began to prevent or suppress forest fires, 

effectively excluding fire from the landscape for nearly 80 

years (Aldrich and others 2010, Flatley and others 2013). 

Long-term exclusion of fire has led to major changes in 

forest structure, function, and composition, particularly 

among forest types dominated by yellow pines and oaks. 

For example, excluding fire has increased the density of 

fire-sensitive trees and shrubs, which, in turn, have 

prevented pine and oak regeneration, shaded out grasses 

and forbs, and reduced the diversity of vegetation across 

the Southern Appalachians (Harrod and others 1998, 2000; 

Turrill and others 1995).

Since the mid-1990s, land managers throughout the 

Appalachians have sought to use natural and prescribed 

fires to reverse the effects of fire exclusion. Fire exclusion, 

however, has contributed to a buildup of wildland fuels 

that make wildland fires more difficult to control, and 

that pose a threat to forest health: when these forests 

eventually burn, they often burn with undesirable 

intensity and/or severity (Reilly and others 2012; Vose 

2000, 2003). As a result, land managers restoring fire 

in the Southern Appalachians face two interrelated 

questions: first, how to effectively reduce hazardous fuels, 

and second, how to restore fire-dependent communities, 

especially pine, pine and oak, and oak forests, while 

minimizing undesirable effects.

Hazardous fuels

Wildland fuels in Appalachian forests fall into two general 

categories—live and dead. Live fuels consist primarily of 

evergreen shrubs, particularly mountain laurel, that can 

pose serious problems for fire control, but do not typically 

contribute significantly to available fuels during landscapelevel burns. Dead fuels, on the other hand, are flammable 

vegetation at or near the forest surface, such as leaf litter, 

duff, and woody debris. Organic duff is the most common 

form of dead fuel (50-70 percent of the total). Other dead 

fuels include litter (10-20 percent) and logs >3 inches in 

diameter (also called 1,000-hour fuels, 10-20 percent). 

These fuel classes are not consumed at the same rate by 

dormant-season burning (Jenkins and others 2011, Vose 

and others 1999, Waldrop and others 2010). Dormantseasons burns, which occur in late winter and early spring, 

consume relatively high amounts of litter, but most of the 

heavier, longer-burning fuels are not consumed.

In contrast, Jenkins and others (2011) found late summer 

and fall burns consumed a much higher percentage of 

duff and 1,000-hour fuels. These growing-season burns 

generally coincided with the annual peak of the drought 

index for the region (as measured by the Keetch-Byram 

Drought Index; Keetch and Byram 1968). Although higher 

levels of heavy fuel consumption were associated with 

successful pine regeneration, they were also strongly 

correlated with higher levels of mortality in the pine and 

oak overstory, which led to large increases in fuel loading 

as dead trees fell to the ground. In addition, growingseason burns and wildland fires frequently increase the 

rate at which nonnative plants invade the community (see 

Kuppinger 2008).
11

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

Pine and oak restoration

Specific objectives for restoring pine and oak communities 

usually center on reducing the abundance of fire-sensitive 

trees and shrubs, increasing pine and oak regeneration, 

and increasing the abundance of grasses and forbs. Several 

burning techniques have been used to achieve these 

objectives, with mixed results.

Single, and even multiple, low-intensity burns (backing/

flanking fires with flame length <3 feet) during the 

dormant season have not achieved objectives for pine and 

oak restoration (Chiang and others 2005, Elliott and Vose 

2005a, Jenkins and others 2011). In general, pine and/or 

oak regeneration did not increase following low-intensity 

burns, and, although all of the studies documented initial 

reductions in fire-sensitive trees and shrubs, these and 

other studies also documented prolific and repeated basal 

resprouting for many of these species.

High-intensity burns (headfires with flame length >8 feet) 

have also been used during the dormant/early season to 

address pine and oak restoration objectives. A common 

response to high-intensity, early-season burns, which has 

not been widely reported, is for these fires to kill large 

numbers of overstory trees, creating large, stand-level gaps 

that subsequently become dominated by hardwood resprouts. 

This can happen with fires at any time of the year, although 

pines can regenerate after late-season fires where a seed 

source exists (Jenkins and others 2011). High-intensity 

burns have been shown to be successful in regenerating 

Table Mountain pine (Waldrop and Brose 1999), and may 

contribute to oak regeneration in formerly pine-dominated 

sites (Elliott and others 2009). In general, however, these 

types of fires are not effective in regenerating oak stands, 

and are not recommended for restoration projects, due to 

concerns about fire control, burn effectiveness, and the loss 

of seed trees (Brose and others 2006, Elliott and others 2009, 

Jenkins and others 2011, Waldrop and Brose 1999).

In summary, we have found the combination of vegetation 

change and fuel accumulation attributed to fire exclusion, 

coupled with the topographic complexity of the landscape 

and the operational constraints in applying fire, poses 

a conundrum for land managers in the Southern 

Appalachians.

Burning too hot—Burning too hot (high-intensity or 

dry-season burning) may reduce high levels of fuels and 

establish pine regeneration, but will typically produce 

undesirable levels of overstory mortality, which, in turn, 

will increase forest fuel loading, decrease the oldest 

structural components in the stand, exacerbate the loss of 

seed trees, increase the growth of undesirable hardwood 

resprouts through overstory release, and facilitate the 

invasion of nonnative plant species.

Burning too cool—Burning too cool (low-intensity or 

dormant-season burning) may avoid some of the negative 

effects of high-intensity burning, but could produce lessthan-desired fire spread, fuel reduction, and reductions 

in fire-sensitive trees and shrubs. As a result, restoration 

objectives such as pine regeneration may not be met in 

a timely fashion, especially while seed trees continue to 

diminish across the landscape.

A fire restoration strategy

What, then, is the best course of action for reintroducing 

fire into long-unburned sites to restore oak and pine 

communities in the Southern Appalachians? Because the 

consequences of burning too hot are far more difficult to 

overcome than the consequences of burning too cool, we 

believe the most cautious approach is to begin with cool, 

dormant-season burns, and to gradually increase burn 

severity by varying burn season and fire intensity. Our 

experience in the Great Smoky Mountains has led us to a 

multiple-burn strategy with the following features:

l Frequent burning (2- to 7-year intervals) over a 20-year 

period.

l Gradual reduction of heavier fuels such as duff and 

coarse woody debris.

l First- and second-entry burns that are primarily low 

intensity (flame length <3 feet) and occur in the dormant 

or very early growing season. The goal for first- and 

second-entry burns is to avoid creating stand-level (>2 

acre) canopy gaps that increase fuel loads and release 

fire-sensitive resprouts. This is most important during 

first entry.

l Subsequent burns that increasingly use variable 

intensity and seasonality to reduce fuels and achieve 

desired community structure and composition.

l Some form of monitoring that can be the basis for 

adaptive management. Monitoring can include simple 

visual assessments, burn severity maps, or various types 

of plots.

Restoring fire to long-unburned sites is a long-term 

process that will require the insight of fire researchers, 

as well as the experience and skill of fire managers. The 

strategy outlined above is, at best, an informal consensus, 

and may not be applicable to the broad range of regional 

fire management objectives or site-specific conditions. 

Additionally, this management approach does not consider 

the use of mechanical treatments, which may contribute 

to the rapid creation of desired forest structure, but which 

are typically limited to small-scale projects. Rather, this 

strategy constitutes a general set of guidelines that should 
12

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

allow managers to achieve long-term vegetation and 

fuels objectives across broad landscapes while avoiding 

negative outcomes. The most important principles 

embedded in this strategy are the reliance on moderation, 

patience, and adaptive management in the application of 

varied combinations of fire intensity and seasonality to 

meet regional goals for fuels management and pine/oak 

restoration.

— Rob Klein, Great Smoky Mountains National Park, 

Gatlinburg, TN

What are the consequences if we don’t use 1.9 fire?

Fire was widespread and frequent throughout much of 

the Eastern United States both before and after European 

settlement (Abrams 1992, Fowler and Konopik 2007, Van 

Lear and Waldrop 1989). Beginning in the 1920s, however, 

fire was actively suppressed, changing plant communities 

across the region (Clark 1990, Wolf 2004). In oak and pine 

communities, these changes combined to produce dense 

forests dominated by mesophytic tree species.

Changes in community structure—Fire suppression 

changed the structure of many plant communities, as open, 

fire-dependent communities such as prairies and savannas 

were invaded by woody species (Abrams and Nowacki 

1992). Compared to presuppression communities, oak-pine 

communities are now structurally dense, with stem densities 

as much as 10 times higher (Nowacki and Abrams 2008). 

Higher tree densities have increased stand basal areas, 

despite declines in average tree diameters, because the stands 

contain many more trees in smaller size classes (Fralish and 

others 1991).

Changes in community composition—In the absence 

of fire, mesophytic tree species, such as yellow poplar, 

maple, and cherry, tend to be competitively superior to 

more xeric oak and pine species. Fire suppression allowed 

fire-sensitive, shade-tolerant mesophytic species to replace 

more fire-dependent, shade-intolerant oaks and pines 

(Nowacki and Abrams 2008).

Changes in community composition affect the future role 

of fire in the community, because increases in mesophytic 

tree species decrease the likelihood of fire (Abrams 1992, 

Nowacki and Abrams 2008). For example, the high leaf 

area of shade-tolerant, mesophytic species casts heavy 

shade and limits air movement, decreasing wind speeds, 

increasing relative humidity, and creating a moist, cool 

forest floor (Nauertz and others 2004). Shady, moist 

conditions reduce understory flammability both directly, 

because the community retains moisture more effectively, 

and indirectly, because the additional moisture promotes 

the decomposition of forest fuels.

Changes in fuel loads—Fire suppression also changed 

the fuels in oak and pine communities (Washburn and 

Arthur 2003). Compared to the leaves of mesophytic 

trees, oak leaves are typically thicker, stiffer, and more 

resistant to decomposition (Abrams 1990, Carreiro and 

others 2000). Their rigid and irregular structure allows 

oak leaves to dry more effectively, and remain dry over a 

longer period of time, than mesophytic leaves, improving 

aeration, and therefore flammability, in the litter layer 

(Scarff and Westoby 2006). Mesophytic leaves, on 

the other hand, tend to lie flat and adhere to the forest 

floor, trapping moisture, minimizing air pockets, and 

enhancing decomposition (Lorimer 1985, Van Lear 2004). 

Oak leaves also contain high amounts of lignin, which 

delays decomposition, allowing oak leaves to remain in 

the litter for a relatively long time (Cromack and Monk 

1975). The leaf litter produced by mesophytic tree species 

tends to contain small amounts of lignin, and the leaves 

decompose rapidly into a moist organic layer that is more 

likely to resist burning (Nowacki and Abrams 2008, 

Washburn and Arthur 2003).

Fuel loads are also determined by the amount of woody 

debris in the stand. In general, woody debris that is dry and 

retained in the community for a long time increases the 

flammability of the stand (Nowacki and Abrams 2008). The 

decomposition rates of woody debris, however, follow the 

rates of leaves, with oak and hickory debris decaying at the 

slowest rates, followed by beech, then maple (MacMillan 

1988). Tyrrell and Crow (1994) reported that oak logs (with 

a half-life of 40 years) and pines (13-16 years) were more 

resistant to decay than mesophytic species such as maple 

(6-15 years).

All of these changes—increases in stand density, shifts in 

community composition, and decreases in fuel loads—

reduce the flammability of oak and pine communities in 

the Southern Appalachians. This process—fire suppression 

leading to increases in mesophytic species that, in turn, 

reduce the flammability of the community—has been called 

mesophication (Nowacki and Abrams 2008). It appears to 

be a common outcome in oak and pine forests wherever 

fire has been suppressed (Bond and others 2005). The more 

mesic and fertile the ecosystem, the more rapidly it will 

undergo mesophication (Nowacki and Abrams 2008).

Once communities become mesophytic, however, returning 

fire and fire-adapted communities to the landscape can 

be challenging, due to the increased difficulty of burning, 

the loss of fire-adapted species, and the increased costs 

associated with the restoration (Abrams 2005). As a 
13

THE ROLE OF FIRE IN THE SOUTHERN APPALACHIANS

result, the mesophication of Southern Appalachian forests, 

especially oak-pine forests, is likely to continue (Nowacki 

and Abrams 2008).

In summary, fire suppression, especially in oak-pine 

and pine communities, has produced structural and 

compositional changes in Southern Appalachian forests 

that have led to a more mesophytic condition. This socalled mesophication of oak and pine forests becomes a 

positive feedback loop because mesophytic trees species 

produce leaf litter and woody debris that is less likely to 

burn than oaks and pines, further suppressing fire. In the 

absence of prescribed fire, we expect mesophication of 

oak and pine forests to continue, increasing the challenges 

facing land managers as they attempt to restore oak and 

pine forests in the Southern Appalachians.

— Tom Waldrop, Center for Forest Disturbance Science, 

Clemson, SC

— Duke Rankin, USDA Forest Service, Southern Region, 

Atlanta, GA
14

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

Succession is an orderly progression of changes in 

community characteristics following disturbance, from 

early successional communities characterized by open 

conditions and ruderal species, to late successional 

communities characterized by closed conditions and 

competitive species (Odom and Barrett 2005, Odum 1969). 

As plant communities undergo succession, so do wildlife 

communities: some wildlife requires early successional 

communities for habitat, while other wildlife requires midor late successional habitats.

Ecologists often distinguish between primary succession, 

on sites previously unoccupied by vegetation, and 

secondary succession, on sites previously occupied by 

vegetation. Secondary succession typically follows some 

form of disturbance to the existing plant community, and 

the nature of the disturbance influences the composition, 

structure, and trajectory of successional change over time. 

In the Eastern United States, secondary succession has 

traditionally been associated with abandoned agricultural 

fields (e.g., Cadenasso and others 2008). Because these 

abandoned fields had been cleared of native vegetation, 

they exhibited dramatic changes in both species 

composition and community structure as they underwent 

succession to native forest. In the forested communities of 

the Southern Appalachians, however, succession is often 

initiated by active management and forest regeneration. 

Because managed forests regenerate largely via stump 

sprouts and localized seedlings, the composition of a 

regenerating forest is usually quite similar to the original 

forest. In this context, the term “succession” is frequently 

used to describe changes in community structure.

As a result, recent studies of both succession and 

early successional habitat have focused on functional 

characteristics of successional communities. Lorimer (2001), 

for example, distinguishes between early successional 

habitat, which he defines as communities dominated by 

species characteristic of highly disturbed areas, and young 

forest habitat, which he defines as young stands of later 

successional, forest species. More recently, Greenberg and 

others (2011a) have used the term early successional habitat 

to describe a wide variety of communities characterized by 

(1) an absence of a closed, mature tree canopy; (2) a welldeveloped ground or shrub component; and (3) young trees.

In the context of forest management and restoration in 

the Southern Appalachians, early successional habitat is 

primarily a wildlife resource, produced by either timber 

harvests or active management, such as the construction 

of wildlife openings. Compared to undisturbed forests, 

2. EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS

regenerating forests share many characteristics with 

traditional forms of early successional habitat, such as 

small, rapidly growing trees and understory species 

associated with open conditions. As a result, we believe 

young, regenerating forests can be functionally equivalent 

to more traditional forms of early successional habitat, 

especially for many wildlife species, and especially during 

the first several years following harvest.

For the purposes of this paper, we will use the wildlife 

definition of early successional habitat, typically 

considered a combination of regenerating forests and 

wildlife openings. When considered a wildlife resource, 

early successional habitat may also be produced by natural 

disturbances, such as wildland fire, windstorms, or the 

natural formation of canopy gaps.

In researching these questions, Southern Research 

Station scientists identified a need for a symposium 

on early successional habitat in the Southern 

Appalachians. A 1-day symposium on was held on 

April 8, 2010, as part of the annual meeting of the 

Association of Southeastern Biologists in Asheville, 

North Carolina. This symposium addressed the 

ongoing decline of many plants and animals 

associated with early successional habitat in the 

Eastern Upland Hardwood Forest region. Presenters 

also synthesized the current knowledge about early 

successional forests and the wildlife associated 

with these habitats, further addressing many of the 

questions posed in this paper. A contributed volume 

that includes proceedings of the symposium, edited by 

Cathryn H. Greenberg, Beverly Collins, and Frank R. 

Thompson III, was published in summer 2011 and is 

available online at www.springer.com/life+sciences/

ecology/book/978-94-007-1619-3.

Why are early successional forests 2.1 important habitats for wildlife species?

Early successional forests provide two critical resources for 

wildlife: habitat structure and cover, and forest foods.

Habitat Structure and Cover

Vegetation structure, measured both as the vertical 

strata within a forest and as the distribution of forest 

types across the landscape, has a strong impact on the 

diversity and composition of wildlife communities 

(MacArthur and MacArthur 1961, Thatcher and others 
15

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

2007). Disturbance can enhance biological diversity, 

especially at the landscape scale, by creating a mosaic 

of habitats or successional stages (see Franzreb and 

others 2011, Shifley and Thompson 2011). Many 

wildlife species in the Southern Appalachians use early 

successional habitats to meet various biological needs, 

including foraging, hunting, nesting, rearing young, 

escape, thermoregulation, and protection from the 

elements (Dickson 2001). Other species use a variety of 

successional stages, but require early successional habitats 

during a particular biological season or time of year. 

Some wildlife species do not require early successional 

habitats, but are more abundant in these habitats, and, in 

general, their populations and individuals are healthier 

when a variety of successional stages are available (Fuller 

and DeStephano 2003).

Birds—Many bird species characteristic of early 

successional habitat in the Southern Appalachians are 

declining as abandoned pastures and farmlands return to 

forest and existing forests mature (Askins 2001, Franzreb 

and others 2011, Shifley and Thompson 2011). Several 

studies report higher bird species richness, diversity, 

and density in sites that were disturbed by management 

activities (Annand and Thompson 1997, Baker and Lacki 

1997) or natural disturbance (Blake and Hoppes 1986, 

Greenberg and Lanham 2001) compared to mature, 

undisturbed forest.

Bird species associated with early successional habitat 

include eastern bluebird, American goldfinch, chestnutsided warbler, golden-winged warbler, yellow-breasted chat, 

eastern meadowlark, field sparrow, northern bobwhite, 

loggerhead shrike, and indigo bunting. Different species, 

however, are associated with different types of early 

successional habitat. For example, meadowlarks are found in 

and around herbaceous openings relatively clear of woody 

encroachment (Roseberry and Klimstra 1970), while goldenwinged warblers use herbaceous openings with considerable 

shrub/bramble encroachment (Klaus and Buehler 2001, 

Litvaitis 2001). When these areas succeed into dense thickets 

and brush, yellow-breasted chats and brown thrashers will 

be present (Burhans and Thompson 1999, Stauffer and Best 

1980). Other species, such as chestnut-sided warblers, are 

found primarily in very young forest (<8 years) with high 

stem densities (King and Byers 2002), while ruffed grouse 

are found primarily in young forest (6-20 years; Jones and 

others 2008, Tirpak and others 2010).

Other bird species use early successional habitat 

periodically during the year. For example, wild turkeys 

in the Southern Appalachians preferentially use openings 

dominated by forbs for brooding (McCord and Harper 

2011). American woodcocks perform courtship rituals 

in grassy openings during the winter, but forage in 

young forest stands inside riparian zones (Dessecker and 

McAuley 2001). Cerulean warblers usually nest in mature 

closed-canopy stands, but forage in canopy gaps (Weakland 

and Wood 2005). Many bird species associated with mature 

forest commonly bring their fledglings to early successional 

forest, presumably because these habitats offer high-quality 

foraging as well as dense, protective cover (Greenberg and 

Lanham 2001, Whitehead 2003).

Mammals—Mammals associated with early successional 

habitat in the Southern Appalachians include the 

Appalachian cottontail, hispid cotton rat, and groundhog. 

White-tailed deer commonly use herbaceous openings 

with shrubby cover, which may influence fawn survival 

(Beier and McCullough 1990, Piccolo and others 2010). 

Young forests are an important source of deer browse 

during spring and summer for approximately 7 years after 

regeneration harvest (Johnson and others 1995). Canopy 

closure, however, eventually reduces browse in these 

regenerating forests (Johnson and others 1995), and the 

stands are not important food resources for deer until hard 

mast becomes available. Black bears will also frequent 

openings and young forest stands during summer when 

soft mast is available. The availability of early successional 

habitat may reduce bear movements and has been linked 

to smaller home ranges (Litvaitis 2001). Several species 

of eastern bats increase foraging activity in recently 

harvested stands (Loeb and O’Keefe 2011, Tichenell and 

others 2011).

Herpetofauna—Salamanders that require cool, moist 

conditions generally decline in clearcuts because of 

their increased risk of desiccation, and populations may 

take years to recover (Petranka and others 1993, 1994; 

Tilghman and others 2012). On the other hand, Raybuck 

(2011), working in western North Carolina, did not detect 

a significant change in salamander abundance after 

shelterwood harvests or other silvicultural treatments 

designed to promote oak regeneration, such as prescribed 

fire. Some reptile populations, such as fence lizards, 

increase in recently disturbed areas, likely because of 

improved opportunity for thermoregulation and foraging 

(Greenberg 2001, Moorman and others 2011, Russell and 

others 2004).

Forest Foods

Young, upland hardwood forests also function as highquality food patches by providing abundant fruit, nutritious 

foliage, and flowers that attract pollinating and foliar 

arthropods, which, in turn, support high populations of 

small mammals. These small mammals and arthropods 

become prey for numerous vertebrate predators. In the 

Southern Appalachians, many species of native birds and 

other vertebrates forage opportunistically in young stands 

for fruit and arthropods.
16

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

Fleshy Fruit—Native fleshy fruit is a key food resource 

for both game and nongame wildlife (Martin and others 

1951). Most birds and mammals consume fruit at least 

occasionally (Martin and others 1951, Willson 1986), 

and fruit can be a critical resource for both migratory 

birds in the fall (Willson 1986) as well as resident and 

overwintering birds in the winter, when arthropods and 

other forest food sources are scarce (Borgmann and 

others 2004, Greenberg and Forrest 2003, Kwit and 

others 2004, McCarty and others 2002, Whitehead 2003). 

Fruit consumption has also been linked to mammalian 

survival and reproductive success (Eiler and others 1989, 

Rogers 1976).

Compared to closed-canopy conditions, fleshy fruit 

production is much greater in forest openings caused 

by either natural disturbance (e.g., Blake and Hoppes 

1986, Thompson and Willson 1978) or by silvicultural 

disturbance such as harvesting (e.g., Greenberg and 

others 2007a, Mitchell and Powell 2003, Perry and others 

1999). Fruit production in the Southern Appalachians 

closely correlates with stand age and achieves its highest 

levels during the first 10 years following two-age harvest 

(Greenberg and others 2011b). Greenberg and others 

(2007a), for example, found dry pulp biomass of fleshy 

fruit was similar in young two-age stands and mature 

forest during the first 2 years following harvest, but by 

year three fruit production was 5.0 to 19.6 times higher in 

young stands compared to mature forest. Canopy openings 

facilitate the establishment of disturbance-mediated species 

such as pokeweed and blackberry, both of which produce 

large amounts of fleshy fruits, and increases in available 

light, moisture, and space provide optimal growing 

conditions for many fruit-bearing plants, including species 

typically associated with mature forests that may already 

be present in the disturbed areas. For example, huckleberry 

is a prolific fruit producer in mature forests and recovers 

rapidly after harvest (Greenberg and others 2007a, Powell 

and Seaman 1990). Stump sprouts of dogwood, American 

holly, Fraser magnolia, black cherry, sassafras, and 

blackgum also produce fruit 1-3 years following harvest. 

As a result, land managers can enhance fruit availability 

for many game and nongame species by creating or 

maintaining patches of young forest (see Greenberg and 

others 2007a, 2011b).

Arthropods—Arthropods play important ecological 

roles as predators and prey (Hammond and Miller 1998), 

decomposers (Moldenke and Lattin 1990), nutrient cyclers 

(Asquith and others 1990), herbivores (Wilson 1987), and 

pollinators (Westman 1990). They also represent a large 

proportion of biological diversity and support invertebrate 

and vertebrate diversity by serving as an important 

food resource. In the Southern Appalachians, leaf litterdwelling arthropods are often positively correlated with 

leaf litter depth, moisture, or both (Duguay and others 

2000, Greenberg and Forrest 2003, Harper and others 

2001, Haskell 2000, Whitehead 2003). In general, litterdwelling arthropods are more abundant and compose more 

biomass in mature forests, where shade and thick leaf litter 

provide a cooler, moister microclimate compared to young 

forest. Flying and foliar arthropods, however, are more 

abundant in young stands where tree density and canopy 

cover are low (Whitehead 2003), most likely due to higher 

concentrations of young, palatable foliage on shrubs, tree 

sprouts, and herbaceous plants. Pollinating insects tend to 

be most abundant in open, disturbed conditions, such as 

clearcuts (Healy 1985), recently burned forests with dead 

trees (Campbell and others 2007), roadsides planted with 

clover or orchard grass (Hollifield and Dimmick 1995), and 

herbaceous wildlife openings (Harper and others 2001), 

because these areas contains the highest concentrations of 

flowering plants (Greenberg and others 2011b).

Forage—Forage, which includes herbaceous plants and 

woody browse, is important for several wildlife species, 

including white-tailed deer, cottontails, black bears, 

wild turkeys, and ruffed grouse. Forage is most readily 

available in constructed wildlife openings, which can 

provide 1,000-10,000 pounds of forage per acre per 

month (Harper 2008). The amount of forage in forests, 

on the other hand, is influenced by the amount of light 

penetrating the forest canopy (Ford and others 1993; see 

Greenberg and others 2011b). Closed-canopy forests in the 

Southern Appalachians typically provide 50-150 pounds 

of dry weight forage per acre within 4.5 feet of the ground, 

while recently harvested stands provide approximately 

1,000 pounds of dry weight forage per acre (Beck and 

Harlow 1981, Della-Bianca and Beck 1985). Available 

forage declines significantly 5-7 years after harvest 

because the developing canopy closes the opening (Beck 

and Harlow 1981, Johnson and others 1995).

Management practices that allow at least 30 percent light 

to penetrate the canopy also increase forage availability. 

For example, shelterwood harvests will typically double 

or triple forage availability compared to closed-canopy 

conditions, and, when followed by low-intensity prescribed 

fire, may provide a sevenfold increase in forage availability 

(Jackson and others 2007, Lashley and others 2011, Shaw 

and others 2010).

— Katie Greenberg, Bent Creek Experimental Forest, 

Asheville, NC

— Craig Harper, University of Tennessee, Knoxville, TN
17

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

What species are favored by early 2.2 successional habitat and how strong is 

the relationship?

Many species of birds, mammals, and reptiles use early 

successional habitat in the Southern Appalachians. Some 

species use openings seasonally, or for specific needs such 

as foraging. Other species use these areas year-round, or 

for multiple needs, such as nesting, escape, foraging, and 

courtship.

Birds—Historically, almost the entire Appalachian region 

was forested, with disturbed areas created by natural factors 

such as canopy gaps, fires, insect infestations, grazing 

by native species, and localized weather conditions such 

as hurricanes, tornadoes, and ice storms. Askins (1995), 

however, argues that native grassland and other successional 

habitats were an integral part of the pre-European landscape. 

Native American agricultural practices are believed to 

have profoundly affected the landscape, primarily from 

slash-and-burn methods that cleared forest for agricultural 

use. As crops depleted soil nutrients, Native Americans 

abandoned old cropland and cleared additional lands, 

creating a presettlement landscape consisting of forests and 

fields in varying stages of succession (Askins 1999, Patterson 

and Sassaman 1988). These abandoned croplands would 

have provided critical habitat for disturbance-dependent 

species. In addition, forest understory was burned by Native 

Americans to improve conditions for hunting small game 

and deer (Van Lear and Harlow 2000).

Many disturbance-dependent bird species are associated 

with more than one type of successional community, and 

may use a combination of communities such as savannas, 

woodlands, gaps in mature forest, and shrubby cover. In 

Eastern North America, 128 bird species considered rare, 

or believed to be declining in much of their range, are 

associated with these communities (Hunter and others 

2001). Rare or declining species that use disturbancemaintained habitats include the Appalachian yellow-bellied 

sapsucker, eastern and Appalachian Bewick’s wren, goldenwinged warbler, cerulean warbler, Swainson’s warbler, and 

indigo bunting.

Due to recent large-scale reductions in the amount and 

distribution of early successional habitat, most disturbancedependent birds have experienced a decrease in population 

numbers, which is expected to continue (Franzreb and 

others 2011, Hunter and others 2001, Shifley and Thompson 

2011). According to Breeding Bird Survey data from 

1966 to 1994 for species associated with shrub cover, 10 

species (58.8 percent) in the Ridge and Valley, 8 species 

(50.0 percent) in the Cumberland Plateau, and 9 species 

(61.5 percent) in the Blue Ridge Mountains Physiographic 

Regions appeared to have undergone significant 

population declines (Franzreb and Rosenberg 1997). Early 

successional habitat may also be important to species 

usually associated with mature forest (Pagen and others 

2000) because some species may seek patches of disturbed 

habitat at least during part of their life cycle (Anders and 

others 1998, Vega Rivera and others 1998).

Mammals—Early successional habitat, particularly gap 

openings, can be important foraging areas for several bats 

in the Southern Appalachians (Loeb and O’Keefe 2011). 

Larger species such as the hoary bat, silver-haired bat, big 

brown bat, and eastern red bat often forage in areas with 

reduced clutter, such as wildlife openings, small cut areas, 

and gaps within intact forest (Krusic and others 1996, Loeb 

and O’Keefe 2006, Owen and others 2004). Even some 

smaller bats such as pipistrelles frequently use openings 

(Loeb and O’Keefe 2006, Schirmacher and others 2007). 

The northern long-eared bat, however, avoids openings 

and prefers closed-canopy forests, presumably because its 

echolocation and flight characteristics allow the species 

to forage effectively in forests (Loeb and O’Keefe 2006, 

Patriquin and Barclay 2003).

In contrast, early successional communities are rarely used 

for roosting. Male eastern red bats occasionally roost in 

early successional habitat in the Southern Appalachians 

(O’Keefe and others 2009), but most bats in the Southern 

Appalachians roost in trees in mature forests during the 

summer and hibernate in caves or migrate south during 

the winter. Early successional forests, therefore, are not 

important habitats for bat roosting ecology. Because early 

successional forests are an important foraging habitat for 

some species, however, the presence of these forests may 

influence roost site selection. For example, male eastern 

pipistrelles in the Southern Appalachians roost closer than 

expected to small, nonlinear openings and two-age harvest 

areas <5 years old, and red bats roost closer than expected 

to linear openings such as gated roads and trails (O’Keefe 

and others 2009). As a result, small openings scattered 

among mature forests may be beneficial to many bat 

species in the Southern Appalachians.

Mammals other than bats appear to be less obligated 

to early successional forests. Litvaitis (2001) suggests 

that a few mammals are habitat specialists (e.g., eastern 

cottontails). Some carnivores (e.g., bobcats) rely on early 

successional habitat for prey items, and a few other 

mammals (e.g., black bears) depend seasonally on the 

abundant fruit found in many forest openings.

Reptiles—Canopy removal results in higher light levels; 

warmer, drier microclimates; and reduced leaf litter cover 

on the forest floor. These conditions, especially warmer 

temperatures, appear to benefit reptiles (Adams and others 

1996, Greenberg 2002, Phelps and Lancia 1995; see also 
18

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

Moorman and others 2011). Most reptile species require 

warm temperatures for egg incubation and the successful 

development of hatchlings (Deeming and Ferguson 1991, 

Goin and Goin 1971). Hotter, drier microclimates can 

also facilitate movement and thermoregulation for many 

reptile species. In the Southern Appalachians, lizards in 

general, and fence lizards in particular, may become more 

common in sites with reduced canopy cover, including 

clearcuts (McLeod and Gates 1998) and large canopy 

gaps (Greenberg 2001). Lizards may also become more 

common after one (Greenberg and Waldrop 2008) or two 

(Matthews and others 2009) prescribed burns.

— Kay Franzreb, Bent Creek Experimental Forest, 

Asheville, NC

— Katie Greenberg, Bent Creek Experimental Forest, 

Asheville, NC

— Susan Loeb, Clemson University, Clemson, SC

Are some wildlife species negatively affected 2.3 by creating early successional habitat?

Several studies in the eastern hardwood forest suggest 

that heavy canopy removal treatments such as clearcuts or 

shelterwoods can adversely affect the local abundance of 

terrestrial salamanders (Ash 1988, 1997; deMaynadier and 

Hunter 1995; Harpole and Haas 1999; Petranka and others 

1993, 1994; Pough and others 1987; Reichenbach and Sattler 

2007; Russell and others 1999, 2004). Canopy removal 

produces higher light levels; warmer, drier microclimates; 

and reduced leaf litter cover, which can desiccate 

salamanders (Crawford and Semlitsch 2008a, 2008b; 

deMaynadier and Hunter 1995; Renken 2006; Russell and 

others 2004). Following clearcutting, salamanders and 

frogs can retreat underground for short periods, evacuate 

the harvest area, or perish (Semlitsch and others 2008), 

and declines in salamander populations may not be fully 

realized for 2-3 years following clearcutting (Ash 1988). 

In the Southern Appalachians, salamanders may virtually 

disappear from sites following clearcutting, and may not 

return to preharvest levels for 20 years (Ash 1988, 1997; 

Petranka and others 1993, 1994; but see also Adams and 

others 1996; Harper and Guynn 1999). Recovery time for 

salamanders tends to vary with leaf litter, which, in turn, 

appears positively correlated with community moisture 

levels—mesic forests recover faster than dry forests 

(Moorman and others 2011). Full recovery, however, may 

take many years. Crawford and Semlitsch (2008b), for 

example, found stream salamanders and their terrestrial 

habitat were less common in Southern Appalachian forest 

stands <40 years of age compared to stands >41 years of 

age. Tadpoles of some frog species may develop faster or 

survive better in ponds inside clearcuts (Semlitsch and 

others 2009), but juvenile and adult stages still require 

forested habitat that may require many years to recover.

On the other hand, Raybuck (2011), working in western 

North Carolina, did not detect a significant change in 

terrestrial salamander abundance after shelterwood 

harvests. Frogs and toads do not appear to be strongly 

affected by forest disturbance (Moorman and others 2011), 

and disturbances that retain heavy canopy cover, such as 

midstory removal, two-age harvests, selection harvests, 

firewood cutting, thinning, heavy browsing by deer, and 

low-intensity burns, do not affect terrestrial salamander 

abundance (Adams and others 1996, Brooks 1999, Floyd 

2003, Ford and others 1999, Greenberg and Waldrop 2008, 

Harpole and Haas 1999, Homyack and Haas 2009, Knapp 

and others 2003, Matthews and others 2009, Messere and 

Ducey 1998, Moseley and others 2003, Pough and others 

1987). Small harvest area and connections to high-quality 

habitat can also mitigate impacts and provide refuge and 

recolonization sources.

Harvest effects on stream-breeding salamanders may also 

be mitigated by the use of riparian buffer zones. Peterman 

and Semlitsch (2009) found that larval salamanders in 

streams were negatively impacted by 30-foot riparian 

buffers, likely due to increased stream sedimentation, and 

suggested 100-foot buffers to reduce the effects of timber 

harvest on larval salamanders in Southern Appalachian 

streams. Crawford and Semlitsch (2007) recommended a 

300-foot streamside buffer zone to provide core terrestrial 

habitat for stream-breeding salamander species in the 

Southern Appalachians, many of which use terrestrial, 

riparian habitat during much of their adult lives.

— Katie Greenberg, Bent Creek Experimental Forest, 

Asheville, NC

Do tree species differ in their response to 2.4 early successional habitat?

Although tree species most likely form a gradient in 

their response to canopy disturbance, canopy trees are 

traditionally separated by foresters into three groups: 

species tolerant of overstory shade, species somewhat 

tolerant of overstory shade, and species intolerant of 

overstory shade (Baker 1949). Tolerant species, such 

as beech and eastern hemlock, tend to grow slowly, but 

persistently, under closed-canopy conditions. Intolerant 

species, on the other hand, include many desirable timber 

and wildlife species, such as red oak and yellow-poplar. 

Compared to tolerant species, intolerant species grow 
19

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

vigorously in open conditions, but tend to grow very poorly 

beneath closed canopies and are unlikely to persist without 

some form of canopy disturbance. Species of intermediate 

tolerance, such as black cherry and white pine, are able 

to grow slowly under partial shade. Although species in 

all three groups may respond positively to gap formation, 

open conditions created by forest management, such as 

timber harvest, generally favor intolerant tree species at the 

expense of tolerant tree species.

This basic concept, however, can be influenced by a variety 

of factors, most notably by the preharvest composition 

of a stand. Following clearcut or two-age harvest of a 

mature forest in the Southern Appalachians, the initial 

composition of the regenerating stand is largely a reflection 

of the preharvest stand, especially on dry sites. With few 

exceptions, most Appalachian hardwood species with 

commercial value are intolerant of shade and require welldeveloped root systems to begin, and maintain, the height 

growth necessary to achieve and hold a canopy position. 

In addition, most hardwood species produce prolific basal 

sprouts when the main stem is cut, and utilize nutrients 

stored in their roots to quickly initiate height growth. As 

a result, tree species that are not present in the stand at the 

time of the harvest do not have an adequate seed source, 

are unable to grow quickly following germination, and are 

unlikely to become established in the regenerating stand.

On moist sites, canopy composition of the postharvest 

stand at crown closure often differs from the preharvest 

stand because rapid height growth from some mesophytic 

species, particularly yellow-poplar and red maple, are 

capable of producing intense competition (Beck and 

Hooper 1986). In addition, the ability of seedlings to grow 

quickly in height following release from competition, 

especially for seedlings of yellow-poplar and sweet 

birch, is another important characteristic that partially 

determines the eventual composition of early successional 

forests (Loftis 1990a). As a result, both shade tolerance 

and the growth response of tree species in the preharvest 

stand must be considered when predicting the outcome of 

silvicultural prescriptions.

Tree species may also differ in their response to open 

conditions created through natural disturbances. Generally, 

gap phase reproduction correlates poorly with the 

reproduction of tree species (Bray 1956), although at least 

two studies have reported significant correlations between 

gap size and tree reproduction. Rankin and Tramer (2002a), 

working in the Appalachian province of southeastern 

Ohio, reported significant correlations between gap size 

and the tree species successfully colonizing former canopy 

gaps. Dale and others (1995), working in hardwood forests 

in the Midwest, reported shade-tolerant species were 

more frequent in smaller openings, and the proportion 

of intolerant species increased as the size of the opening 

increased. In addition, oaks and hickories were more 

common in openings on the poorer sites, while yellowpoplar was more common in openings on the better sites.

— W. Henry McNab, Bent Creek Experimental Forest, 

Asheville, NC

— David Loftis, Bent Creek Experimental Forest, 

Asheville, NC

— Duke Rankin, USDA Forest Service, Southern Region, 

Atlanta, GA

How much early successional habitat is 2.5 necessary to provide for biodiversity? What 

is the optimal percentage of early successional 

habitat to meet desired goals?

The answer to this question depends primarily on the scale 

at which biodiversity is measured. Biodiversity occurs at 

several levels: at the community level, along a gradient 

between communities, and at the landscape or regional level 

(Noss 1983, see also Whittaker 1972). Early successional 

habitat would most likely enhance biodiversity along 

gradients between communities (Sharitz and others 1992).

Presumably, the optimal amount of early successional 

habitat for a variety of wildlife species varies between the 

6-9 percent of the forest that would occur in natural gapphase disturbance at any given moment (Runkle 1982) and 

the 60 percent of the forest estimated by Lorimer (2001) at 

peak disturbance levels during the late 19th century.

Although the natural gap disturbance rate would seem to be 

the minimum necessary to provide for biodiversity, Hunter 

and others (2001) argue that natural disturbance regimes, 

especially gap-phase disturbances now largely absent from 

a landscape with few old-growth forests, are insufficient to 

maintain disturbance-dependent bird species. DeGraaf and 

Yamasaki (2003) suggest that to optimize early successional 

species diversity, early successional habitat should compose 

10-20 percent of the forest landscape, or roughly twice the 

natural gap rate. This recommendation is very close to 

the 10- to 20-percent estimate of naturally occurring early 

successional habitat derived in Question 2.6, below.

Biodiversity, however, does not vary directly with early 

successional habitat (Shifley and Thompson 2011). Most of 

the plant diversity in Southern Appalachian forests occurs 

in the herb layer, which varies across several environmental 

gradients, including disturbance. In the Southern 

Appalachians, herbaceous diversity responds positively 
20

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

to overstory disturbance by increasing the percentage of 

shade-intolerant species in the herbaceous layer (Shifley 

and Thompson 2011). Shade-tolerant herbs also respond 

positively to at least some overstory disturbance, but the 

response is presumably brief and confined to a small 

window of time immediately following the disturbance, 

before shade-intolerant herbs colonize and dominate the 

site (see Rankin and Tramer 2000b). When overstory 

disturbance is severe, forest herbs may take decades to 

recover to pre-disturbance levels (Elliott and others 1997, 

2011). Once lost, many forest herbs are severely limited by 

seed dispersal, and may take many years to recolonize a 

forest stand (Elliott and others 2011).

— Duke Rankin, USDA Forest Service, Southern Region, 

Atlanta, GA

What is the natural distribution of early 2.6 successional habitat across the landscape?

For the purposes of this report, we define natural 

disturbance in the Southern Appalachians as any process 

creating early successional habitat that is not explicitly 

due to human actions. Using this definition, natural 

disturbances would include natural fires, windstorms, 

hurricanes, landslides, outbreaks of native insects, and 

autogenic forest dynamics, such as gap fall. This definition 

would exclude modern silvicultural actions, such as timber 

harvests and prescribed burning. Although we recognize the 

potential role of Native Americans in shaping the Southern 

Appalachian landscape, especially through the use of fire, 

we consider these affects anthropogenic in nature and 

exclude them from our definition of natural disturbances.

Evidence on the amount and distribution of natural 

disturbance are derived from several sources, including 

historical records, old-growth forest stands, pollen and 

charcoal deposits, and direct observation of contemporary 

disturbances (Lorimer 2001). While each of these lines 

of evidence can provide important information, each also 

has its limitations, and quantitative estimates of natural 

disturbance must be interpreted cautiously. Lorimer 

(2001) lists some of the challenges for determining natural 

disturbance patterns:

l Because of the long history of human occupation in 

the Southern Appalachians, evidence on natural and 

anthropogenic disturbance is easily entangled.

l Changes in global climate may alter disturbance 

regimes over geological time.

l Due to soil, topographic, and human settlement 

patterns, disturbance patterns on the landscape are 

spatially nonrandom.

l Although potentially long-lasting, severe disturbance 

is highly episodic and spatially heterogeneous, making 

estimates of its impacts problematic.

Because disturbance patterns depend upon biotic and 

abiotic factors that differ at regional scales, disturbance 

patterns also differ by geographic region (Runkle 1990). In 

the Southern Appalachians, natural disturbance patterns 

are often attributed primarily to gap-phase dynamics (e.g., 

Runkle 1981). Gap-phase disturbance in eastern deciduous 

forests averages about 1 percent of the landscape per year 

(Runkle 1982). Because canopy gaps remain in early 

successional habitat for approximately 6-8 years following 

formation, approximately 9 percent of the landscape is in 

some form of gap-phase reproduction at any time (Rankin 

and Tramer 2002a). Gap size varies by the number of trees 

involved in the disturbance, but, in the Appalachian region, 

generally ranges between 1,000 and 5,000 square feet 

(Rankin and Tramer 2002a).

Gap-phase disturbance, however, tends to occur in small, 

localized patches that retain forest characteristics (Runkle 

and Yetter 1987) and is generally not considered equivalent 

to more-extensive disturbances created by catastrophic 

events such as windstorms, infestations, and wildland 

fire (Shure and others 2006). For example, gap-phase 

disturbances tend to produce small openings that typically 

close more quickly than more extensive disturbances that 

create larger openings. Although qualitatively different, 

more-extensive disturbances are also characteristic of the 

Southern Appalachians, and may be just as important as 

gap-phase disturbance.

Windstorms—Greenberg and McNab (1998), using data 

compiled by Neumann and others (1993), report hurricanerelated windstorms have recurred in the Southern 

Appalachians 14 times since 1871, at intervals ranging from 

1 to 24 years. Hurricane Opal damaged approximately 0.3 

percent of the national forest lands in northeastern Georgia, 

western North Carolina, and eastern Tennessee in 1995. 

Although Greenberg and McNab (1998) calculate that 

approximately 9.3 percent of the landscape would suffer 

some form of windthrow disturbance over the 200-year 

lifespan of many eastern forest trees, only a small portion 

of this time would be spent in the early successional phase. 

If this phase extends 20 years from the initial disturbance, 

then the amount of young forest produced by catastrophic 

windthrow would affect approximately 1 percent of the 

Southern Appalachian landscape at any given time.
21

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

Infestations—Infestations of native insects, such as the 

southern pine beetle, can be widespread and produce 

relatively large patches of young, regenerating forest. The 

U.S. Department of Agriculture, Forest Health Protection 

estimates the total amount of beetle infestation on a yearly 

basis. The estimates are not specific to the Southern 

Appalachians and are presumably based on partial activity 

within forest stands. For 2001, a year characterized by a high 

level of beetle activity, the total area of beetle infestations, 

estimated as one-half of the reported acres for Georgia, 

Tennessee, and North Carolina, is 1.6 million acres, or 

approximately 2-5 percent of the Southern Appalachian 

landscape.

Other insects also affect the Southern Appalachians, 

but many of these species, such as the hemlock woolly 

adelgid and the emerald ash borer, are invasive exotics, and 

presumably would not have contributed to the more natural, 

presettlement disturbance regime. Because these impacts 

have not been fully analyzed on a regional basis, they 

will not be considered further in this report. On the other 

hand, we recognize these species will have an extensive 

impact on the Southern Appalachians and may need to be 

considered in future analyses for early successional habitat.

Fire—High-intensity, catastrophic fire is rare in the 

hardwood forests of the Southern Appalachians. Despite 

extensive research over the past few decades and 

widespread acceptance that fire has been an integral part 

of the Southern Appalachian landscape over both historic 

and prehistoric periods, the amount of early successional 

habitat created by wildland fire cannot be adequately 

characterized. Guyette and others (2006) estimate return 

intervals for presettlement (about 1650-1850) eastern 

deciduous forest in the Southern Appalachians at around 

10 years, although they caution that return intervals are 

highly variable in both time and space. The fire literature 

contains no quantitative estimates for historical levels of 

fire extent and severity in the Southern Appalachians, and 

no estimates regarding the amount of early successional 

habitat produced by fire. If we estimate the amount of early 

successional habitat produced by wildland fire as roughly 

the equivalent of the habitat produced by windstorms 

and insect infestations combined, the total amount of 

habitat present at any given moment would constitute 

between 3 and 6 percent of the landscape of the Southern 

Appalachians.

Ice storms—Ice storms occur regularly in the Southern 

Appalachians (Runkle 1985), with return intervals estimated 

around 20 years (Abell 1934). These storms can damage and 

kill overstory trees, producing canopy openings (Boerner 

and others 1988). Working in southwestern Virginia, 

Warrillow and Mou (1999) estimated a single ice storm 

damaged roughly 0.7 percent of a 1,200-acre forest, although 

localized storm damage can be much higher (Boerner 

and others 1988). Compared to other forms of landscape 

disturbance in the Southern Appalachians, however, the 

effects of ice storms are more variable, and therefore more 

difficult to categorize, especially in relationship to early 

successional habitat. If half the canopy damage caused by 

an ice storm produces early successional habitat, and if the 

early successional habitat remains for 10 years following 

the storm, then early successional habitat produced by ice 

storm damage may occupy around 1 percent of the Southern 

Appalachian landscape at any given moment.

Other disturbances—Other disturbances, such as 

landslides and floods, may produce locally severe 

disturbances. Compared to landscape-scale disturbances, 

such as gap-phase disturbance and hurricanes, however, the 

amount of early successional habitat created by landslides 

and floods appears negligible and therefore will not be 

considered further.

Compared to gap disturbances, catastrophic disturbances 

occur at longer intervals, are highly episodic, and vary 

markedly in time and space (Lorimer 2001, Greenberg 

and others 2011a). As a result, the relative influence of 

gap and catastrophic disturbances at a specific location 

is not always clear. Gap disturbance appears ubiquitous 

in mature, old-growth forest, and, if estimated at 8-10 

percent of the original forest, would be the dominant 

form of disturbance in the mesic forests of the Southern 

Appalachians, especially in protected coves that are 

not prone to windstorms and severe fires. Catastrophic 

disturbance also appears ubiquitous in the Southern 

Appalachians, and, if estimated at 6-12 percent of the 

original forest, would be the dominant form of disturbance 

in more-exposed forests, such as ridges and dry slopes. 

Combining the two estimates, potentially 20 percent of 

the landscape would be in early successional habitat at 

any given time, and roughly half of the early successional 

habitat would be created by catastrophic events.

— Duke Rankin, USDA Forest Service, Southern Region, 

Atlanta, GA

Is there a threshold value for early 2.7 successional habitat patch size?

Threshold values for early successional patch sizes vary 

by the species associated with the habitat. In addition, 

threshold values may depend on the context of the habitat—

for example, the availability of nearby roosting or foraging 

habitat in mature forest may affect the species of bats found 

in early successional openings (Loeb and O’Keefe 2006). 

As a result, threshold values are closely related to both the 
22

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

species under consideration and the context of the habitat 

in the landscape. Quantifying these thresholds may be very 

difficult.

Plants—Rankin and Tramer (2002a, 2002b) used current 

and reconstructed canopy gaps in eastern hemlock forests in 

the Appalachian region of southeastern Ohio to examine the 

role gap size plays in the establishment of both overstory trees 

and understory herbs. They concluded that canopy gaps less 

than approximately 1,000-1,500 square feet play virtually no 

role in the establishment of trees, but gaps >1,500 square feet 

are strongly associated with tree recruitment. This association 

is connected with gap size: tolerant trees, such as hemlock 

and beech, are more likely to become established in mediumsized gaps, while intolerant trees, such as yellow-poplar, are 

confined to the largest gaps. This finding suggests at least 

some tree species in the Southern Appalachians exhibit 

different thresholds for gap size before they can successfully 

reproduce in natural canopy gaps. Understory plants, on 

the other hand, all respond to canopy gaps to some degree, 

but some species appear to utilize sub-gap disturbances to 

persist beneath closed canopies, suggesting that at least some 

understory herbs do not exhibit a lower threshold value for 

canopy disturbance.

Rankin and Tramer (2002a, 2002b) primarily addressed 

lower threshold values for a few species in the Southern 

Appalachians. Although we know of no study that 

explicitly addresses the possibility of upper threshold 

values for Southern Appalachian herbs, these values 

may be correlated with the minimum thresholds for 

the reproduction of shade-intolerant trees, because, at 

that point, the dominant flora presumably changes from 

species characteristic of forest environments to species 

characteristic of open environments. In the Northeastern 

United States, this threshold is roughly 2 acres of open 

conditions (DeGraaf and Yamasaki 2003).

Wildlife—Threshold values of early successional habitat 

for wildlife vary by species. Home ranges for early 

successional species vary from 3 acres for yellow-breasted 

chat (Thompson and Nolan 1973) to as much as 200 acres 

of young forest for male ruffed grouse (Thompson and 

Fritzell 1989). Many wildlife species, however, use early 

successional habitat as a portion of their home ranges, 

making threshold levels difficult to determine. Greenberg 

(2001) and Greenberg and Lanham (2001) suggest that gaps 

as small as 0.25 to 3 acres can provide important habitat for 

some early successional birds and reptiles. Based on their 

studies in the Northeast, DeGraaf and Yamasaki (2003) 

recommended group selection harvests of at least 2 acres, 

although small clearcut openings (20-30 acres) appear to 

meet the habitat needs of more bird species than group 

selection openings (Costello and others 2000). White-tailed 

deer will use small group selection harvests—typically, 

openings between 0.25 and 0.5 acres—for foraging and 

bedding, but larger harvests may be necessary to regenerate 

trees in the presence of high deer densities. Well-distributed 

harvests of approximately 20 acres have been recommended 

to avoid problems with tree regeneration due to deer grazing 

(Harlow and Downing 1969). Other studies have shown the 

number of bird species increases with increasing size of 

clearcuts, up to 50 acres (Rudnicky and Hunter 1993).

Single-tree selection—the harvest procedure that seems 

most similar to natural canopy gaps—does not create 

openings large enough to provide early successional 

habitats for birds (DeGraaf and Yamasaki 2003). For 

example, golden-winged warblers, one of the early 

successional species of most concern in the Southern 

Appalachians, tend to avoid patches smaller than 5 acres; 

their numbers tend to increase when patch sizes are 

between 30 and 100 acres (Buehler and others, unpublished 

data cited by Hunter and others 2001).

Landscapes—The most problematic values for early 

successional patch sizes occur at the landscape scale: 

How much of a forest, or a watershed, should be in early 

successional habitat? Again, this value depends upon 

the resource associated with the landscape. For example, 

black bears, the wildlife resource with the largest home 

range in Eastern North America, require a mixture of 

successional stages, using both mature forest and early 

successional areas during different times of the year. Even 

for bears, however, neither the optimal nor the threshold 

values of early successional habitat have been determined. 

Working in New England on Northern Appalachian forests, 

DeGraaf and Yamasaki (2003) suggest that, to optimize 

early successional species diversity, early successional 

habitat should compose about 10-20 percent of the forest 

landscape.

— Duke Rankin, USDA Forest Service, Southern Region, 

Atlanta, GA

— Nancy Herbert, Southern Research Station, 

Asheville, NC

How do we sustain early successional 2.8 habitat in the landscape?

By definition, early successional habitat is transient and 

disappears from the landscape unless constantly produced 

by natural or anthropogenic means (Shifley and Thompson 

2011). Since reaching peak levels of 60 percent in the late 

19th century, young forest in North America has declined 

steadily, to ≤20 percent in many regions (Lorimer (2001). 

Due to shifts in land management, declining timber 
23

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

harvests, fire suppression, and urban sprawl, traditional 

methods for generating and maintaining early successional 

habitat—primarily logging, wildland fires, firewood 

cutting, and farm abandonment—are no longer available 

(Trani and others 2001). Early successional habitat is now 

created mostly through active management, including 

timber harvest, fire, and the creation of grass and forb 

wildlife openings (Harper 2007, Litvaitis 2001).

Working in the northern hardwood forests characteristic 

of the Northeastern United States, DeGraaf and Yamasaki 

(2003) offer several suggestions for sustaining early 

successional habitat in managed forests that might apply to 

Southern Appalachian forests, while maintaining a balance 

between early and late successional habitat:

l Use regeneration cuts >2 acres to create favorable 

habitat for the reproduction of shade-intolerant and 

intermediate-tolerant tree species. Harvest areas 

> 2 acres produce dense regenerating stands with an 

abundance of fleshy fruits important for many wildlife 

species. Clustering smaller cuts can also maintain 

adequate amounts of early successional habitat while 

not fragmenting larger patches of older forests, and may 

encourage wider use by a range of wildlife.

l Shorten the time between periodic regeneration cuts 

in a management area to every 10 years. This action 

would produce permanent stands of forest ≤10 years old, 

ensuring their presence in the landscape.

l Because they are maintained in an early successional 

condition, powerline rights-of-way can help meet the 

needs of wildlife associated with early successional 

habitat (King and Byers 2002). Bulluck and Buehler 

(2006), however, caution that powerlines are structurally 

and functionally different from regenerating forest, 

and, while creating habitat for many bird species such 

as Kentucky warblers, powerlines would not support 

bird communities similar to those found in other forms 

of early successional habitat. Powerlines may increase 

the number of bird species, but may also increase the 

number of brood parasites, creating ecological traps 

or sinks (Lanham and Whitehead 2011). Other than 

their effects on birds and butterflies, however, the 

potential values of powerline rights-of-way as early 

successional habitats have been little studied (Lanham 

and Whitehead 2011).

Thompson and DeGraaf (2001) remind us, however, that 

even though some early successional wildlife species seem 

to be in decline, the issue remains controversial. Managing 

for early successional habitat may reduce habitat for at least 

some late successional species, forcing managers to 

make decisions between competing resources. In addition, 

benchmarks and threshold values for early successional 

habitat are not always clear or widely accepted. Finally, 

public land management agencies provide only a small 

proportion of the available young forests in the Eastern 

United States (Trani and others 2001).

— Nancy Herbert, Southern Research Station, 

Asheville, NC

Are natural disturbances such as canopy 2.9 gaps sufficient to maintain early 

successional habitat?

Measured by aerial extent, the rate of gap disturbance 

in mature forests in the Eastern United States is 

approximately 1 percent per year, producing a landscape 

in which 6–9 percent of the forest communities are 

in active canopy gaps (Runkle 1982; see also Rankin 

and Tramer 2002a). Gap disturbance, however, may be 

qualitatively different from disturbance produced by timber 

harvest, and is certainly different from early successional 

habitat created through fire and wildlife openings. Gap 

formation rates in contemporary forests may be less than 

rates in primary forests, because managed forests tend 

to be younger, and therefore less prone to gap formation, 

than fully mature forests (Hunter and others 2001). In 

addition, canopy gaps are often too small to reach the 

thresholds of early successional habitat needed by many 

disturbance-dependent wildlife species (DeGraaf and 

Yamasaki 2003), although both the effective gap size and 

the likelihood of producing early successional habitat may 

be increased through multiple disturbances (Frelich and 

Reich 1999, Rankin and Tramer 2002a). Because of these 

differences, several authors argue that natural processes 

such as gap disturbance can no longer provide sufficient 

early successional habitats (Askins 2000, DeGraaf and 

Yamasaki 2003, Hunter and others 2001).As discussed in 

Question 2.6, however, gap disturbance is only one part of 

the natural disturbance regime. Other disturbances, such 

as wildland fire and windstorms, also produce forms of 

early successional habitat. Compared to gap disturbances, 

however, large-scale disturbances are both less frequent and 

less evenly spread across the landscape. Clearly, large-scale 

disturbances can produce substantial amounts of early 

successional habitat, but these effects tend to be localized, 

temporary and unpredictable, and do not address habitat 

needs across the full range of the Southern Appalachian 

landscape (DeGraaf and Yamasaki 2003).

— Duke Rankin, USDA Forest Service, Southern Region, 

Atlanta, GA
24

EARLY SUCCESSIONAL HABITAT IN THE SOUTHERN APPALACHIANS 

Are openings created by fire sufficient to 2.10 produce early successional habitat?

Yes, fire-created openings are sufficient to produce early 

successional habitat if the fire is sufficiently intense to 

cause stand replacement (Smith 2000). Fire disturbance, 

however, may be qualitatively different from early 

successional habitat produced by timber harvest or natural 

disturbances. Stand-replacing fire, for example, may leave 

large numbers of snags, increasing populations of woodboring insects, woodpeckers, insect- and seed-eating birds, 

and secondary-cavity nesters. Stand-replacing fire is also 

more likely to produce early successional habitat in large, 

contiguous blocks.

— Duke Rankin, USDA Forest Service, Southern Region, 

Atlanta, GA

What are the consequences if we do not 2.11 manage for early successional habitat?

The hardwood forests in the Eastern United States 

are largely a legacy of stand-replacing, anthropogenic 

disturbance between 1700 and the early 1900s. This 

widespread disturbance produced historically high levels 

of early successional habitat (Lorimer 2001). As current 

forests mature, the amount of early successional habitat on 

public lands will most likely decline from these historical 

highs. This decrease, in turn, will probably reduce the 

population sizes, and frequencies, of species that depend 

on this habitat. As a result, species most closely associated 

with early successional habitat, especially golden-winged 

warbler, yellow-breasted chat, ruffed grouse, and American 

woodcock, will likely decrease (Dessecker and McAuley 

2001, Thompson and DeGraaf 2001).

The effects of decreased early successional habitat on 

mammals are not as clear, and few significant effects 

have been documented (e.g., Heske 1995, Menzel and 

others 1999). Litvaitis (2001) argues that some game 

animals, such as turkey, white-tailed deer, ruffed grouse, 

black bear, and bobcat, would be negatively affected by a 

significant reduction in the amount of early successional 

habitat.

On the other hand, older forests will become more prone 

to natural disturbances, especially canopy disturbances 

associated with the fall of large, mature trees. When 

gaps involve multiple trees, or occur in conjunction with 

weather events, the canopy openings may be large enough 

to support early successional habitat (Rankin and Tramer 

2002a). Under this scenario, mature, functional forests 

will eventually produce a shifting equilibrium of early 

successional habitat, proportional to the gap formation rate 

characteristic of the ecosystem.

— Duke Rankin, USDA Forest Service, Southern Region, 

Atlanta, GA
25

OAK REGENERATION IN THE SOUTHERN APPALACHIANS 

Upland, mixed-oak forests occupy >50 percent of the 

forested land in the Central Hardwood Region of the 

United States (Johnson and others 2002), and oak trees 

play a pivotal role in the forest ecology and economy 

of the region. Oaks have been called a keystone species 

in Southern Appalachian forests (Fralish 2004, Petich 

and others 2002), influencing a wide range of species. 

For example, oaks have been associated with the 

distribution, abundance, and behavior of many forms 

of wildlife, ranging from migratory birds to black bear 

(Clark 2004, McShea and Healy 2002, Rodewald 2003). 

Acorns influence small mammal populations that form an 

important prey base for raptors and carnivores (McShea 

2000). And both the physiognomy and decomposition 

rates of oak leaves, as well as the texture of oak bark, can 

increase arthropod abundance and availability for many 

bird species (e.g., Rodewald 2003), contributing to higher 

bird diversity in oak-dominated forests than in mapledominated forests.

In addition to their ecological importance, oaks are among 

the most economically valuable hardwood species (Guyette 

and others 2004). Oaks provide raw materials for a wide 

range of wood products including veneer, lumber, pallets, 

pulp, and paper (Patterson 2004). The wood products sector 

accounts for an estimated 1.9 percent of all jobs and 2.3 

percent of gross regional production across the mainly rural 

South (Abt and others 2002).

Despite the ecological and economic importance of oakdominated forests, their sustainability is now threatened 

by a combination of oak decline, which causes progressive 

oak crown dieback and mortality (Oak and others 2004), 

and widespread oak regeneration failure. This failure is 

generally attributed to changes in disturbance regimes, 

especially fire, which have produced corresponding shifts 

in species composition, increasing the competitive pressure 

on oaks (Aldrich and others 2005). Maintaining oak forests 

in the face of these threats will require active management 

throughout the Southern Appalachians.

What are the current management practices 3.1 for regenerating oak forests?

For upland hardwood stands in the Southern Appalachians, 

forest management is typically focused on restoring 

forest structure, function, and species composition, 

with a particular emphasis on regenerating oak species. 

On the other hand, upland hardwood stands are often 

compositionally complex, dominated by a mixture of oak 

and hickory species in association with hardwood species 

such as maple, yellow-poplar, magnolia, ash, and black 

cherry. Many of these species tend to be mid-tolerant 

of shade, and exhibit a reproductive strategy based on 

a combination of advance regeneration and overstory 

disturbance. For mid-tolerant hardwoods in general, 

and for oaks in particular, successful regeneration is a 

multistep process that includes seed production, seedling 

establishment, seedling development into stems that can 

compete following overstory release, and the release itself 

(Johnson and others 2002). In addition, some moderate 

level of overstory or midstory disturbance that increases 

light levels to the lower levels of the stand is required for 

seedlings to develop into the larger stems necessary to form 

an effective advance reproduction (Loftis 1990b).

The success of oak regeneration is therefore dependent 

upon the size and abundance of advanced oak reproduction 

prior to any silvicultural manipulation (Loftis 1990a, 

Sander 1971, 1972). In mesic oak stands in the Southern 

Appalachians, however, adequate advance reproduction of 

oaks may be uncommon because, although oaks remain 

in the overstory, the midstory is often dominated by more 

shade-tolerant species that suppress oak seedlings. As a 

result, the advance reproduction is composed of smallstemmed plants, and these plants are unable to compete 

effectively following overstory disturbance. Under these 

conditions, regeneration treatments that substantially 

reduce the forest canopy do not regenerate oaks, because 

shade-intolerant competitors, such as yellow-poplar, 

quickly overtop the small-stemmed advanced oak 

regeneration (Loftis 1990b, Shure and others 2006).

When large-stemmed advance reproduction is absent, the 

oak shelterwood method is an effective treatment to ensure 

successful oak regeneration (Loftis 1990b). This system 

can be described as a two-step shelterwood. The first 

cut, also called an establishment cut is a noncommercial 

removal of approximately 20-30 percent of the stand 

basal area from below, using herbicides to reduce resprouting. This effectively removes the shade-tolerant 

midstory, increasing light levels at the forest floor and 

encouraging the development of large-stemmed advanced 

oak reproduction without creating a light environment 

conducive to the establishment and growth of yellowpoplar. Once the oak seedlings have grown into largestemmed advanced (>4 feet in height) reproduction, the 

overstory can be removed—a period of approximately 

10 years. Although initially developed to improve oak 

regeneration on moderate to highly productive stands in 

3. OAK REGENERATION IN THE SOUTHERN APPALACHIANS
26

OAK REGENERATION IN THE SOUTHERN APPALACHIANS 

the Southern Appalachians, the oak shelterwood method 

may also be used to regenerate any species of upland 

hardwoods that relies primarily on large-stemmed advance 

reproduction (Loftis 1983a).

On xeric oak sites, several factors, including lower soil 

moisture, fewer competing tree species, lower stand 

density, and a greater proportion of light reaching the forest 

floor, generally favor oaks (Johnson and others 2002). 

Because oaks are more competitive under these conditions, 

xeric oak stands are more likely to have abundant 

large-stemmed, advanced oak reproduction, and many 

regeneration methods can be used successfully on xeric oak 

stands (Johnson and others 2002).

— Tara Keyser, Southern Research Station, Asheville, NC

In the absence of management, can oaks 3.2 trees regenerate naturally?

Historically, disturbance events such as low-intensity 

surface fires, timber harvesting, logging-associated 

wildland fires, grazing, land-clearing for agriculture, 

and the loss of American chestnut promoted forest 

conditions conducive to the establishment, development, 

and recruitment of oaks in upland hardwood forests 

(Abrams 1992, Lorimer 1993). Because these conditions 

were widespread during the past two to three centuries, 

oaks gained widespread dominance in many forest stands 

throughout the Southern Appalachians.

Although low levels of oak regeneration has been 

observed throughout the Central Hardwood Region, 

local conditions can substantially influence oak 

regeneration. On drier, less productive sites, such as 

ridge tops, oaks may regenerate naturally because the 

tree density and leaf area in the stand are low enough to 

allow sufficient light for oak seedlings to develop into a 

competitive regeneration source. In addition, fast-growing 

competitors, such as yellow-poplar, are typically not 

present in drier stands (Clinton and others 1994, Johnson 

and others 2002). In mesic, more productive sites, such 

as rich coves, oak regeneration is more problematic, 

primarily due to intense competition from fast-growing 

mesophytic species (Johnson and others 2002, Kellison 

1993, Loftis and McGee 1993).

In these mesic sites, successful oak regeneration is a multistep 

process that includes (1) acorn production; (2) germination 

and seedling establishment; (3) seedling development into 

tall (>4 feet), large-stemmed seedlings, creating a pool of 

advance reproduction that can successfully compete with 

faster-growing species; and (4) timely and sufficient release 

of the advance reproduction through some form of overstory 

removal (Johnson and others 2002). For oaks to reliably 

regenerate, all four steps must occur in sequence and within a 

certain timeframe. On mesic, productive sites in the Southern 

Appalachians, poor oak regeneration generally results from 

the failure of the third step—the development of seedlings 

into a competitive advance reproduction. This failure occurs 

because the low-light conditions characteristic of mature 

forest stands decrease both the survival and the growth rate of 

oak seedlings (Loftis 1983a). As a result, mature forest stands 

may contain ample oak saplings, but the seedlings are short 

(<1 foot), small-stemmed, and unable to compete with fastgrowing species following overstory removal (Loftis 1983a, 

1983b).

Because oaks are intermediate in shade tolerance, midstory 

disturbance that increases light levels at the forest floor 

allows oak seedlings to develop into large saplings that 

form an effective advance reproduction (Loftis 1990b). 

When released through overstory removal, this advance 

reproduction is able to effectively compete with fastgrowing competitors, promoting the recruitment of oaks 

into the overstory of the new, developing stand.

In the absence of a midstory disturbance, forest stands 

may develop in one of two ways. If left undisturbed, the 

low-light levels characteristic of mature forests promote 

the growth, and eventual recruitment, of shade-tolerant 

tree species, such as red and sugar maple (e.g., Spetich 

and Parker 1998). If, on the other hand, the overstory is 

removed through harvest or natural disturbance, shadeintolerant tree species, such as the fast-growing yellowpoplar, quickly occupy the site, and eventually dominate 

the stand (Shure and others 2006). As a result, treatments 

that remove the midstory, which promote the development 

of oak seedlings into a competitive advance reproduction 

while preventing the establishment and development 

of competing species, are needed to ensure timely and 

successful oak regeneration (Loftis 1990b).

Because the four steps associated with oak regeneration 

can occur naturally, successful oak regeneration without 

management can, and does, occur at various positions on 

the landscape, but remains unreliable and stochastic at any 

specific location. Although naturally occurring canopy 

gaps may facilitate oak regeneration (e.g., Clinton and 

others 1994), the natural disturbances characteristic of 

Southern Appalachian forests are generally insufficient to 

regenerate oak stands on a consistent and reliable basis, at 

least on mesic, productive sites (Della-Bianca and Beck 

1985, Loftis 2004).

— Tara Keyser, Southern Research Station, Asheville, NC

— David Loftis, Southern Research Station, Asheville, NC
27

OAK REGENERATION IN THE SOUTHERN APPALACHIANS 

Are there treatments that could improve 3.3 forest health in oak communities?

Ultimately, the term “forest health” comes from our human 

perspective. Although we may equate community health 

with productivity and resilience, especially among desired 

species such as oak trees, communities change over time 

through natural processes that are neither good nor bad.

Within this context, the term ”forest health” is typically 

used when considering community changes brought about 

by nonnatural processes, such as nonnative pathogens, 

or changes in the natural disturbance patterns. These 

nonnatural processes tend to diminish the traits we 

value in our native forest communities—for example, 

forest products, native biodiversity, and wildlife values. 

Managing for forest health seeks to reduce the effects 

of these nonnatural processes and to encourage natural 

processes that will enhance the traits we find desirable.

In an oak-hickory forest community, managing for forest 

health typically means managing for oak production and 

regeneration. Managing for oaks may involve regeneration 

harvest, which removes aging trees that become 

increasingly susceptible to oak decline and mortality, 

and creates the open conditions necessary for vigorous 

regrowth of the stand. Other procedures that can improve 

forest health in these stands include improving the timber 

and wildlife attributes through direct management (timber 

stand improvement and wildlife stand improvement, 

guarding against the introduction of exotic pests and 

disease, returning natural disturbance regimes, and 

reducing human impacts, especially to soils and water. 

Fire may play an important role in maintaining healthy oak 

forests, by reducing and regulating insect pests (Ahlgren 

1974, Komarek 1970, Miller 1979).

Early detection of forest health concerns is also an 

important step in reducing impacts to oak trees. One 

potential indicator of forest stress is the ratio of standing 

dead to live trees, sometimes called the forest health 

quotient. The quotient averages 0.08 in healthy Midwestern 

second-growth forests (Spetich and others 1999), and 0.089 

in Arkansas forests (Spetich and Guldin 1999). Forests 

with quotients above these values may indicate forest 

stress. Although this quotient has not been assessed for the 

Southern Appalachians, comparing quotients derived from 

U.S. Forest Service, Forest Inventory and Analysis (FIA) 

data could help identify emerging forest health issues.

— David Loftis, Southern Research Station, 

Asheville, NC

— Marty Spetich, Southern Research Station, Hot 

Springs, AR

What is the role of fire in regenerating oak 3.4 trees?

Prescribed fire is increasingly being used by land managers 

to restore oak communities and promote oak regeneration 

in the Southern Appalachians (Brose and others 2006, Dey 

and Fan 2009, Van Lear and Watt 1993). Fire is typically 

prescribed to reduce competition between fire-tolerant oak 

species and their fire-intolerant competitors, including red 

maple and yellow-poplar, especially in the regeneration 

layer (Waldrop and others 2008), and to increase light 

levels in the understory, producing larger and more 

competitive oak saplings (Barnes and Van Lear 1998).

Fire behavior and intensity, and therefore fire effects, 

vary across the landscape (Albrecht and McCarthy 2006). 

In general, fires burn at lower intensities on lower, more 

mesic slopes than on higher, more xeric slopes. This 

spatial variability in fire behavior and intensity leads to 

differential mortality among tree species, creating different 

types of forests on different sites (Elliott and others 1999). 

For example, prescribed fire can increase the density of 

competing, non-oak species on mesic sites, but it can 

increase the density of small-diameter oak saplings on 

xeric sites (Albrecht and McCarthy 2006).

The differential effect of prescribed fire across the 

landscape, especially across moisture and productivity 

gradients, emphasizes the need for site-specific 

information on the relationship between site quality and 

oak regeneration. For example, in the Cumberland Plateau 

region of Kentucky, both single and repeated fires have 

been effective at reducing overstory stem density and 

basal area, leading to increased light in the stand, but 

these benefits for oak regeneration are quickly offset by 

increased sprouting of competing species such as sassafras 

and red maple (Alexander and others 2008). Although fire 

can reduce the number of small (<1 foot tall) red maples, 

sprouts from larger, topkilled red maple stems often grow 

more quickly than oak sprouts (Alexander and others 

2008), which can accelerate the loss of oaks from the 

community (Wendel and Smith 1986).

One treatment that has proven effective in regenerating 

oak forests in the Piedmont region of Virginia is the 

shelterwood/burn technique proposed by Brose and others 

(1999a, 1999b). In the first step, the stand is heavily cut 

to release all regeneration sources, regardless of species. 

This initial cut—technically, the establishment cut of 

a shelterwood harvest—promotes root growth in oak 

saplings and stimulates yellow-poplar seeds in the litter and 

duff layers to germinate (Brose and others 1999b).

After the establishment cut, stump sprouts from harvested 

trees advance reproduction present in the stand, and 
28

OAK REGENERATION IN THE SOUTHERN APPALACHIANS 

new seedlings all grow and develop under the sparse 

shelterwood canopy. During this period, oaks allocate a 

substantial proportion of their carbohydrates to root growth 

instead of shoot growth (Kolb and others 1990).

Three to 5 years after the establishment cut, the developing 

stand is burned to control species composition. Oak 

saplings, with large carbohydrate reserves in the root 

system and dormant buds located below the soil surface 

(Burns and Honkala 1990), will often re-sprout vigorously 

after being topkilled by the fire, while competing species, 

such as yellow-poplar and red maple, are often incapable of 

sprouting after being topkilled because they have smaller 

carbohydrate reserves in their roots (Kolb and others 1990), 

and hold their dormant buds above the soil surface (Beck 

1981). The differential response to topkill increases the 

number of oaks relative to the competing tree species.

Interestingly, traditional shelterwood methods (e.g., 

Loftis 1983b, Schuler and Miller 1995) or single 

applications of prescribed fire (Johnson 1974, Wendel 

and Smith 1986), when used separately, rarely enhance 

the competitive position of oak regeneration on highquality sites in the Appalachians. Unless present as large 

advance reproduction, oaks are unable to outcompete 

yellow-poplar seedlings that germinate and develop 

after shelterwood harvest (Loftis 1983b, 1999b), and 

prescribed fire alone appears ineffective at creating the 

light conditions necessary for the development of large 

advance reproduction (Alexander and others 2008). In 

addition, prescribed fire appears ineffective at releasing 

and recruiting advance oak reproduction into larger 

size classes (Alexander and others 2008), unless the fire 

intensity is sufficient to cause mortality in the forest 

canopy (Hutchinson and others 2005b, Signell and 

others 2005).

The combination of shelterwood regeneration followed 

by prescribed fire shows promise in regenerating oaks. 

However, even this method has been proven effective only 

on sites of moderate productivity in the Piedmont region of 

Virginia (site index between 70 and 80 feet; base age 50 for 

white oak; Brose and others 1999a). The shelterwood/burn 

technique has not been tested on high productivity sites, 

such as the cove forests of the Southern Appalachians, 

where competition from yellow-poplar is intense and 

oak regeneration is especially problematic. Researchers 

with the Southern Research Station are currently testing 

the shelterwood/burn technique across a range of sites 

in the Southern Appalachians of North Carolina and the 

Cumberland Plateau of Tennessee.

The recent and widespread-use of prescribed fire to 

regenerate and restore upland oak forests remains relatively 

untested across the various physiographic subregions 

of the Southern Appalachians. Most studies suggest 

prescribed fire, on its own, does not substantially improve 

oak regeneration (Alexander and others 2008, Hutchinson 

and others 2005b, Signell and others 2005, Wendel and 

Smith 1986). Because the effects of fire are spatially and 

temporally heterogeneous, more research is needed to 

determine the effects of prescribed fire, either alone or 

in combination with silvicultural treatments, to improve 

and promote oak regeneration in the upland forests of the 

Southern Appalachians.

— Tara Keyser, Southern Research Station, Asheville, NC

— David Loftis, Southern Research Station, Asheville, NC

What are the effects of cutting oak trees on 3.5 high elevation sites?

High elevation sites in the Southern Appalachians are 

generally associated with elevations greater than 4,000 

feet. At these elevations, forests tend to be cooler and 

moister than lower elevation forests, exhibit shorter 

growing seasons, and develop into forest communities 

more characteristic of northern regions. High elevation oak 

forests are separated from other oak forests by increased 

dominance of mesic oaks, especially northern red oak, and 

reduced dominance by white oak and hickories. They are 

also subject to increased levels of air pollution, especially 

acid rain, and are often considered particularly sensitive to 

environmental changes.

High elevation red oak forests rarely undergo Forest 

Service management actions. As a result, little research 

has been conducted on the effects of actions such as 

timber harvest and prescribed fire. In one study, Elliott and 

Knoepp (2005) examined the regeneration of high elevation 

oak forests. Situated at 4,500-5,500 feet, stands dominated 

by northern red oak were harvested using a variety of 

standard techniques, including two-age, shelterwood, and 

group selection. Six years later, northern red oak remained 

a common constituent of the community in the two-age and 

shelterwood units.

Management actions may also affect high elevation soils. 

Compared to low elevation forests, high elevation forests 

receive higher rates of atmospheric nitrogen deposition. 

As a result, high elevation soils contain seven times more 

total nitrogen than soils in low elevation, mixed-oak forests 

(Knoepp and others 2000, 2008). Because the soils contain 

higher levels of nitrogen, nitrogen cycles through the 

ecosystem faster than at lower elevations, (Knoepp and others 

2000, 2008), increasing nitrogen availability in the soil and 

elevating nitrogen concentrations in streams draining high 
29

OAK REGENERATION IN THE SOUTHERN APPALACHIANS 

elevation watersheds (Knoepp and others 2008, Swank and 

Douglass 1975, Swank and Vose 1997). Disturbance increases 

both the rate of nitrogen cycling through the ecosystem and 

the rate at which nitrogen is exported in streams (Swank 

1988). These responses are similar for both management 

actions and natural disturbances, and are proportional to the 

size of the disturbed area.

In summary, timber harvest in high elevation forests 

appears to have little effect on overstory composition. On 

the other hand, all disturbances at high elevation increase 

the rate at which nitrogen is exported from the ecosystem.

— Jennifer Knoepp, Coweeta Hydrologic Laboratory, 

Otto, NC

What are the consequences if we do not plan 3.6 for oak regeneration?

Changes in disturbance regimes—both natural and 

anthropogenic—are promoting the gradual conversion of 

forests dominated by oaks into forests dominated by shadetolerant species such as red maple (Nowacki and Abrams 

2008, Orwing and Abrams 1994), or by shade-intolerant 

species such as yellow-poplar (e.g., Beck and Hooper 1986, 

Rodewald 2003). Over time, a decrease in the abundance of 

oaks, combined with associated changes in forest structure 

and composition, could have cascading effects throughout 

the upland hardwood ecosystems of the Southern 

Appalachians. For example, acorns are considered a critical 

forest resource because of their influence on populations of 

small mammals (McShea 2000). In turn, small mammals 

disperse spores and seeds (Wolff 1996) and represent an 

important prey base for raptors and carnivores. 

In addition to the critical role of oaks for wildlife habitat 

and as a food resource, the loss of oak, in combination with 

concomitant increases in shade-tolerant species such as 

beech and maple, has been shown to dramatically reduce 

herbaceous plant diversity by reducing the amount of light 

at the forest floor and increasing the depth of the leaf litter 

(Fralish 2004). The loss of oak-dominated forests would also 

reduce ecosystem heterogeneity at the landscape level. This 

could, in turn, result in reduced resistance and resilience to 

future disturbances, including climate change, and the loss 

of ecosystem functioning (Yachi and Loreau 1999).

— Tara Keyser, Southern Research Station, Asheville, NC

— David Loftis, Southern Research Station, Asheville, NC
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41

ACKNOWLEDGMENTS / CONTRIBUTING SCIENTISTS

ACKNOWLEDGMENTS 

The editors gratefully acknowledge our authors, who gave 

so generously of their time and expertise. We also thank 

Beverly Collins, Jaclyn Schnurr, Charles Kwit, Craig 

Harper, and William Coates for their reviews and insights 

on early versions of the manuscript. Following Nancy 

Herbert’s retirement, Katie Greenberg served as project 

liaison with the Southern Research Station. Finally, we 

thank the USDA Forest Service and the Southern Research 

Station for their support throughout the project. 

CONTRIBUTING SCIENTISTS

Kathleen (Kay) Franzreb is a Research Wildlife 

Biologist, U.S. Department of Agriculture (USDA) 

Forest Service Southern Research Station, Bent Creek 

Experimental Forest (Asheville, NC), but is stationed with 

the Department of Forestry, Wildlife, and Fisheries, 274 

Ellington Plant Science Building, University of Tennessee, 

Knoxville, TN 37996. She conducts research related to the 

management and conservation of Neotropical migratory 

birds in the Southern Appalachians, focusing on forest 

management. She also examines the ecology of endangered 

and threatened bird species, with a special interest in redcockaded woodpeckers and Florida scrub-jays. She may be 

contacted at 865-974-0932 or kfranzreb@fs.fed.us.

Cathryn (Katie) H. Greenberg is Project Leader 

and Research Ecologist with the Upland Hardwood 

Ecology and Management Research Work Unit, USDA 

Forest Service Southern Research Station, Bent Creek 

Experimental Forest, 1577 Brevard Road, Asheville, NC 

28806. Her research focus includes (1) effects of forest 

management practices (such as timber harvesting, fuel 

reduction practices, and prescribed fire) and natural 

disturbances (such as wind damage) on plant and animal 

communities, and (2) production of forest food resources, 

such as native fleshy fruit and hard mast, in relation to 

forest types and silvicultural disturbances. She can be 

contacted at 828-667-5261 x118 or kgreenberg@fs.fed.us.

Craig A. Harper is a Professor of Wildlife Management 

and the Extension Wildlife Specialist at the University 

of Tennessee, 280 Ellington Plant Sciences Building, 

Knoxville, TN 37996. He maintains an active extension 

and research program related to managing early succession 

as well as forested areas for wildlife. Of particular 

interest is the use of prescribed fire in maintaining 

early succession and the effect of prescribed fire on the 

vegetative community and resulting wildlife habitat in 

upland hardwoods. He can be contacted at 865-974-7436 or 

charper@utk.edu.

Nancy Herbert retired after serving 18 years as an 

Assistant Director for Research at the USDA Forest 

Service, Southern Research Station, 200 W.T. Weaver 

Boulevard, Asheville, NC 28804. Prior to coming to the 

Southern Research Station, Nancy was a Research Wildlife 

Biologist in the Northeast and in California. Her experience 

in managing research programs, especially her work with 

silvicultural, wildlife, and fish research teams, has provided 

her with a broad view of restoration issues in the South. In 

retirement, Nancy plans to actively enjoy the forests of the 

Southern Appalachians and the many benefits they provide. 

Since her retirement, Nancy requests that questions 

regarding this document be directed to Kier Klepzig, 

Assistant Director for Research. He can be reached at the 

address above or at 828-257-4306 or kklepzig@fs.fed.us.

William (Bill) A. Jackson is an Air Resource Specialist 

for the National Forest System, USDA Forest Service, 

National Forests in North Carolina, 160A Zillicoa 

Street, Asheville, NC 28801. The focus of Bill’s work 

is to understand how air quality impacts the natural 

resources on Forest Service lands, and how Forest Service 

management activities impact air quality. His areas of 

expertise include the effects of ground-level ozone on 

forest trees, the influence of atmospheric deposition on 

biogeochemical cycling, and how to predict and monitor 

downwind concentrations of particulate matter from 

prescribed fires. He can be contacted at 828-257-4815 or 

bjackson02@fs.fed.us.

Tara L. Keyser is a Research Forester with the Upland 

Hardwood Ecology and Management Research Work Unit, 

USDA Forest Service Southern Research Station, Bent Creek 

Experimental Forest, 1577 Brevard Road, Asheville, NC 

28806. Her research focus includes (1) predicting structural 

and compositional change of Southern Appalachian Highland 

forests resulting from natural or silvicultural disturbances, 

including prescribed fire, and developing silvicultural 

treatments to control forest structure and composition; (2) 

modeling the effects of intermediate stand management of 

forest stand dynamics, including carbon storage; and (3) 

quantifying climate-tree growth relations for the predominant 

tree species in the Southern Appalachians. She can be 

contacted at 828-667-5261 x102 or tkeyser@fs.fed.us.

Robert (Rob) N. Klein is a Fire Ecologist with the 

National Park Service, Great Smoky Mountains National 

Park, 107 Park Headquarters Road, Gatlinburg, TN 37738. 

He manages a regional fire-effects monitoring program, 

with an emphasis on the role of scientific monitoring in 

adaptive management of prescribed and wildland fire. He 

is active in planning, ecological monitoring, research, and 

prescribed fire implementation in numerous National Park 

Service units, from the Appalachian Mountains to the 

Southeastern Coastal Plain. He can be contacted at 

865-430-4751 or rob_klein@nps.gov.
42

CONTRIBUTING SCIENTISTS

Jennifer Donaldson Knoepp is a Research Soil Scientist, 

USDA Forest Service Southern Research Station, Coweeta 

Hydrologic Laboratory, 3160 Coweeta Lab Road, Otto, 

NC 28763. She conducts research examining the effects 

of forest management, such as cutting or prescribed 

and wildland fire, on nutrient cycling and soil nutrient 

availability and effects on vegetation. She also examines 

long-term forest nutrient cycling patterns and their changes 

due to changes in nitrogen and sulfur deposition patterns as 

well as climate change. She can be contacted at 

828-524-2128 or jknoepp@fs.fed.us.

Susan Loeb is a Research Ecologist, USDA Forest 

Service Southern Research Station, 237 Lehotsky Hall, 

Clemson University, Clemson, SC 29634. She conducts 

research on the ecology and conservation of forest bats 

in the Southeastern United States. Her studies focus on 

the ecology of rare and endangered species, including the 

Indiana bat, the effects of forest management on forest bat 

habitat use, and methods for inventorying and monitoring 

forest bats. She can be contacted at 864-656-4865 or 

sloeb@fs.fed.us.

Since beginning work on this report, David Loftis

has retired from the Upland Hardwood Ecology and 

Management Research Work Unit, Southern Research 

Station, at the Bent Creek Experimental Forest in 

Asheville, NC. His research focused on the ecology and 

management of Southern Appalachian hardwood forests, 

especially predicting and controlling the composition of 

species following forest regeneration. He also worked on 

decision models for the management of hardwood forests, 

and the effects of silvicultural practices on biological 

diversity.

W. Henry McNab is a Research Forester, USDA 

Forest Service Southern Research Station, Bent Creek 

Experimental Forest, 1577 Brevard Road, Asheville, NC 

28806. His research assignments have included silviculture 

of loblolly pine, smoke management from prescribed 

burning, forest biomass estimation, and silviculture of 

upland hardwoods. For the past 20 years, his research has 

involved ecological classification of plant communities in 

the Southern Appalachian Mountains, with an emphasis 

on quantifying the relationships among environmental 

gradients, species composition, and productivity of forest 

sites. His current research includes developing alternatives 

to site index for classification of forest site quality. He can 

be contacted at 828-667-5261 or hmcnab@fs.fed.us.

Roger Ottmar is a Research Forester with the Fire and 

Environmental Research Applications Team, USDA Forest 

Service Pacific Northwest Research Station at the Pacific 

Wildland Fire Sciences Laboratory, 400 North 34th Street, 

Suite 201, Seattle, WA 98103. He has been involved with 

fuels, fire, and smoke-related research for 33 years and has 

authored more than 100 research publications and final 

reports. He can be contacted at 206-732-7826 (office), 

206-849-3172 (cell), or rottmar@fs.fed.us.

W.T. (Duke) Rankin is the Threatened and Endangered 

Species Program Manager at the USDA Forest Service’s 

Southeast Regional Office, 1720 Peachtree Road NW, Suite 

700, Atlanta, GA 30309. He has performed research on the 

secondary succession of Piedmont forests, the impacts of 

fire to tallgrass prairie, and the understory and regeneration 

dynamics of canopy gaps in eastern forests. He is also 

on the graduate faculty of Western Carolina University, 

where he has contributed to research on nonnative invasive 

plants, early successional habitat, and the effectiveness of 

constructed vernal ponds. He can be contacted at 

404-347-3981 or drankin@fs.fed.us.

Martin (Marty) A. Spetich is a Research Forester, 

USDA Forest Service Southern Research Station, Upland 

Hardwood Ecology and Management Research Work Unit, 

P.O. Box 1270, Hot Springs, AR 71902. Marty’s research 

focus is to address practical, applied forest management 

needs through a combination of current scientific thought 

and needs expressed by field practitioners. His integrated 

research program addresses forest dynamics and the 

development of both short- and long-term studies at the 

stand, State, and regional scale. His research has included 

studies of forest restoration, growth, woody species 

reproduction, competitive capacity, stand dynamics, stand 

composition, forest ecology, disturbance ecology, oak 

decline, land use history, landscape ecology, and effects of 

fire on stand development. He can be contacted at 

501-623-1180 x105 or mspetich@fs.fed.us.

Thomas A. Waldrop is Research Forester and Team 

Leader for Fire Science at the Center for Forest Disturbance 

Science, USDA Forest Service, Southern Research 

Station, 239 Lehotsky Hall, Clemson, SC 29634. His 

work emphasizes fire ecology in the Piedmont and 

Southern Appalachian Mountains, with research on fire 

techniques for stand replacement of Table Mountain pine, 

fire history, fuel loading characteristics of the Southern 

Appalachians, and impacts of stand replacement fire on 

various components of mountain ecosystems. In more 

recent years, he has been a leader in the National Fire and 

Fire Surrogate Study, which studies ecological impacts of 

fire and mechanical fuel reduction techniques in numerous 

ecosystems of the United States. He is currently developing 

the Consortium of Appalachian Fire Managers and 

Scientists to enhance science delivery for fire professionals 

in the Central and Southern Appalachian region. He can be 

contacted at 864-656-5054 or twaldrop@fs.fed.us.
43

INDICES

INDICES

Subject Index

Advance oak regeneration Q3.1 Q3.2 Q3.4

Air pollutants Q1.5 Q3.5

Amphibians Q1.3

Bats Q1.3 Q1.7 Q2.1 Q2.2 Q2.7

Bats foraging Q1.3 Q2.2

Bats maternity colonies Q1.3

Bats roosting Q1.3 Q2.2

Biodiversity Q1.6 Q1.7 Q1.8 Q2.1 Q2.5 Q3.3

Birds Q1.3 Q2.1 Q2.2 Q2.5 Q2.7 Q2.8 Q2.10

Canopy gaps Q1.6 Q2.1 Q2.2 Q2.4 Q2.7 Q2.9 Q2.11

Coarse woody debris Q1.2 Q1.4 Q1.6 Q1.7 Q1.8 Q1.9

Deer browse Q2.1 Q2.3 Q2.7

Fire, Anthropogenic Q1.1 Q2.6

Fire, Behavior Q1.2

Fire, Catastrophic Q1.6 Q2.6

Fire, Effects to air quality Q1.5

Fire, Effects on soils Q1.2 Q1.4 Q1.7

Fire, Frequency Q1.1 Q1.6 Q1.7

Fire, Fuels Q1.1 Q1.2 Q1.5 Q1.6 Q1.7 Q1.8 Q1.9

Fire, Historic Q1.1 Q1.8

Fire, Intensity Q1.1 Q1.2 Q1.3 Q1.4 Q1.6 Q1.7 Q1.8 

Q2.1 Q2.3 Q2.6 Q2.10 Q3.2 Q3.4

Fire, Multiple Q1.3 Q1.4 Q1.8 Q2.2 Q3.4

Fire, Objectives Q2.1 Q2.2

Fire, Season Q1.2 Q1.4 Q1.5 Q1.8

Fire, Smoke Q1.5

Fire, Suppression Q1.1 Q1.6 Q1.8 Q1.9

Fire, Types Q1.2

Forest structure Q1.1 Q1.2 Q1.3 Q1.6 Q1.7 Q1.8 Q1.9 

Q2.1 Q3.1 Q3.6

Forest, Acidic Cove Q1.6

Forest, Bottomland Hardwood Q1.6

Forest, Mixed Mesophytic Q1.6

Forest, Northern Hardwood Q1.6

Forest, Oak-Hickory Q1.4 Q3.3

Forest, Rich Cove Q1.6 Q3.2 Q3.3 Q3.4

Forest, Spruce-Fir Q1.6

Forest, White Pine-Hemlock-Hardwood Q1.6

Frogs Q1.3 Q2.3

Gap-phase dynamics Q1.6 Q2.4 Q2.5 Q2.6 Q2.7 

Q2.9 Q2.11

Herbaceous diversity Q1.2 Q1.6 Q1.8 Q2.5 Q2.7 Q3.6

Herpetofauna Q1.3 Q2.1

Ice storms Q1.6 Q2.2 Q2.6 Q2.2 

Litter Q1.3 Q1.5 Q1.6 Q1.7 Q1.8 Q1.9 Q2.1 Q2.2 Q2.3 

Q3.4 Q3.6

Nonnative invasive species Q1.6 Q1.8 Q2.6

Oak forests Q1.1 Q1.4 Q1.6 Q1.8 Q1.9 Q3.3 Q3.4 Q3.5 Q3.6

Pine forests Q1.1 Q1.6 Q1.8 Q1.9

Pollinators Q2.1

Powerlines Q2.8

Reptiles Q1.3 Q2.1 Q2.2 Q2.7

Riparian buffer zones Q2.3

Salamanders Q1.3, Q2.1 Q2.3

Shade tolerance Q2.4 Q2.5 Q2.7 Q2.8 Q3.1 Q3.2 Q3.4 Q3.6

Small mammals Q1.3 Q1.7 Q2.1 Q2.2 Q3.6

Snags Q1.3

Superfog Q1.5

Timber harvest, Clearcut Q2.1 Q2.2 Q2.3 Q2.4 Q2.7

Timber harvest, Firewood Q2.3

Timber harvest, Group Selection Q2.3 Q2.7 Q3.5

Timber harvest, Historic Q1.1

Timber harvest, Regeneration Q2.1 Q2.8 Q3.3 Q3.5

Timber harvest, Shelterwood Q2.3 Q3.1 Q3.4 Q3.5

Timber harvest, Single-tree Q2.7

Timber harvest, Thinning Q1.2 Q1.4 Q1.7 Q2.3

Timber harvest, Two-age Q2.1 Q2.2 Q2.3 Q2.4 Q3.5

Toads Q1.3 Q2.3

Windstorms Q1.6 Q2.2 Q2.6

Woodpeckers Q1.3 Q2.10

Author Index

Franzreb Q2.2

Greenberg Q1.3 Q2.1 Q2.2 Q2.3

Harper Q1.2 Q2.1

Herbert Q2.7 Q2.8

Jackson Q1.5

Keyser Q1.6 Q3.1 Q3.2 Q3.4 Q3.6

Klein Q1.8

Knoepp Q1.1 Q1.2 Q1.4 Q1.7 Q3.5

Loeb Q1.3 Q2.2

Loftis Q2.4 Q3.2 Q3.3 Q3.4 Q3.6

McNab Q1.5 Q2.4

Ottmar Q1.5

Rankin Q1.6 Q1.9 Q2.4 Q2.5 Q2.6 Q2.7 Q2.9 Q2.10 Q2.11

Spetich Q3.3

Waldrop Q1.1 Q1.2 Q1.4 Q1.5 Q1.6 Q1.7 Q1.9
44

INDICES

Taxonomic Index

Adelgid, hemlock woolly, Adelges tsugae, Q1.6 Q2.6

Ash, American, Fraxinus americana, Q3.1

Bat, big brown, Eptesicus fuscus, Q2.2

Bat, eastern red, Lasiurus borealis, Q1.3 Q2.2

Bat, evening, Nycticeius humeralis, Q1.3

Bat, hoary, Lasiurus cinereus, Q2.2

Bat, northern long-eared, Myotis septentrionalis, Q1.3 Q2.2

Bat, silver-haired, Lasionycteris noctivagans, Q2.2

Bat, Indiana, Myotis sodalis, Q1.3

Bear, black, Ursus americanus, Q2.11 Q2.2 Q2.7 Q2.1

Beech, American, Fagus grandiflolia, Q1.6 Q2.4 Q2.7 Q3.6

Birch, sweet, Betula lenta, Q1.6 Q2.4

Birch, yellow, Betula alleghaniensis, Q1.6

Blackberry, Rubus spp., Q2.1

Blackgum, Nyssa sylvatica, Q2.1

Blueberry, Vaccinium spp., Q1.6

Bluebird, eastern, Sialia sialis, Q2.1 Q1.3

Bobcat, Lynx rufus, Q2.11 Q2.2

Bobwhite, northern, Colinus virginianus, Q2.1

Bunting, indigo, Passerina cyanea, Q1.3 Q2.1 Q2.2

Chat, yellow-breasted, Icteria virens, Q2.1 Q2.7 Q2.11

Cherry, black, Prunus serotina, Q1.6 Q1.8 Q2.1 Q2.4 Q3.1

Cottontail, Appalachian, Sylvilagus obscurus, Q2.1

Cottontail, eastern, Sylvilagus floridanus, Q2.2

Deer, white-tailed, Odocoileus virginianus, 

Q1.6 Q2.1 Q2.11

Dogwood, Cornus florida, Q2.1

Fir, Fraser, Abies fraseri, Q1.6

Frogs, Q1.3, Q2.3

Goldfinch, American, Spinus tristis, Q2.1

Groundhog, Marmota monax, Q2.1

Grouse, ruffed, Bonasa umbellus, Q1.6 Q2.1 Q2.7 Q2.11

Hemlock, eastern, Tsuga canadensis, Q2.7 Q2.4

Hickory, Carya spp., Q3.1 Q3.5

Holly, American, Ilex opaca, Q2.1 

Huckleberry, Gaylussacia spp., Q1.6 Q2.1

Kentucky warblers, Oporornis formosus, Q2.8

Kudzu, Pueraria lobata, Q1.6

Laurel, mountain, Kalmia latifolia, Q1.6, Q1.8

Lizard, fence, Sceloporus undulatus, Q2.2

Magnolia, Magnolia spp., Q3.1

Magnolia, Fraser, Magnolia fraseri, Q2.1

Maple, Acer spp., Q3.1

Maple, red, Acer rubrum, Q1.6 Q2.4 Q3.4 Q3.6

Maple, sugar, Acer saccharum, Q1.6 Q1.8

Meadowlark, eastern, Sturnella magna, Q2.1

Mouse, deer, Peromyscus maniculatus, Q1.3

Mouse, white-footed, Peromyscus leucopus, Q1.3

Oak, chestnut, Quercus coccinea, Q1.6

Oak, northern red, Quercus rubra, Q1.6 Q2.4 Q3.5

Oak, scarlet, Quercus coccinea, Q1.6

Oak, white, Quercus alba, Q1.6 Q3.5

Pine, pitch, Pinus rigida, Q1.6

Pine, shortleaf, Pinus echinata, Q1.6

Pine, Table Mountain, Pinus pungens, Q1.6 Q1.8

Pine, Virginia, Pinus virginiana, Q1.6

Pine, white, Pinus strobus, Q2.4

Pipistrelle, Perimyotis subflavus, Q2.2

Pokeweed, Phytolacca americana, Q2.1

Poplar, yellow-, Liriodendron tulipifera,

Q1.6 Q1.8 Q2.4 Q2.7 Q3.1 Q3.2 Q3.4 Q3.6

Privet, Chinese, Ligustrum sinense, Q1.6

Rat, hispid cotton, Sigmodon hispidus, Q2.1 

Rhododendron, rosebay Rhododendron maximum, Q1.6

Rose, multiflora, Rosa multiflora, Q1.6

Salamander, terrestrial, Plethodon spp., Q1.3, Q2.1, Q2.3

Sapsucker, Appalachian yellow-bellied, Sphyrapicus varius

ssp. appalachiensis, Q2.2

Sassafras, Sassafras albidum, Q2.1 Q3.4

Shrike, loggerhead, Lanius ludovicianus, Q2.1

Sparrow, field, Spizella pusilla, Q2.1

Spruce, red, Picea rubens, Q1.6

Thrasher, brown, Toxostoma rufum, Q2.1

Toads, Q1.3, Q2.3

Turkey, Meleagris gallopavo, Q1.6 Q2.1 Q2.11

Warbler, cerulean, Dendroica cerulean, Q2.1 Q2.2

Warbler, chestnut-sided, Dendroica pensylvanica, Q2.1

Warbler, golden-winged, Vermivora chrysoptera, 

Q2.1 Q2.2 Q2.11

Warbler, Swainson’s, Limnothlypis swainsonii, Q2.2

Warbler, hooded, Wilsonia citrina, Q1.3

Warbler, worm-eating, Helmitheros vermivorus, Q1.3

Woodcock, American, Scolopax minor, Q2.1 Q2.11

Woodpecker, (various species), Q1.3, Q2.10

Wren, Appalachian Bewick’s, Thryomanes bewickii altus, 

Q2.2

Wren, eastern Bewick’s, Thryomanes bewickii, Q2.2
45

APPENDIX

Fire Web sites and Search Options

FRAMES—Fire Research and Management 

Exchange System

http://frames.nbii.gov/portal/server.pt

FRAMES—Southern Fire Portal

http://frames.nbii.gov/southernfire

Fire and Environmental Research Applications

www.fs.fed.us/pnw/fera/

International Association of Wildland Fire

www.iawfonline.org/

Joint Fire Science Program

http://jfsp.nifc.gov/

Association for Fire Ecology

www.fireecology.net/

Fire Learning Network

http://tncfire.org/training_usfln.htm

In particular, click on »South Central FLN (Arkansasbased), »Southeast FLN (Coastal Plain-based), »Southern 

Blue Ridge (Southern Appalachians) and »Appalachians 

(Central Appalachians) and »Network Publications

TreeSearch

www.treesearchfs.fed.us

Tall Timbers Research Station

http://ttrs.org/info/fedbintro.htm

Forestry Encyclopedia Network

http://forestencyclopedia.net

Includes the following encyclopedias: Southern Fire 

Science, Southern Appalachian Forest Ecosystems, 

Environmental Threats, and Southern Bioenergy

Fire Effects Information System

www.fs.fed.us/database/feis

Fire Monitoring

FFI (Fire Effects Monitoring Analysis Software)

http://frames.nbii.gov/ffi

Fire Effects Monitoring Handbook

http://www.nps.gov/fire/download/fir_eco_

FEMHandbook2003.pdf

Basis for the 5140 USFS Region 8 Fire Effects 

Monitoring program

FIREMON

www.fs.fed.us/rm/pubs/rmrs_gtr164.html

USFWS Fuel and Fire Effects Monitoring Guide

www.fws.gov/

Measuring and Monitoring Plant Populations

http://www.blm.gov/nstc/library/pdf/MeasAndMon.pdf

Other Resources

Converting metric units to English

www.srs.fs.usda.gov/landowners/convert_metric_units.htm

Fire Management Today (magazine)

http://www.fs.fed.us/fire/fmt/index.html

InterfaceSouth—Southern Center for Wildland UrbanInterface Research and Information

www.interfacesouth.org/

Communicator’s Guide to Wildland Fire

www.nifc.gov/preved/comm_guide/wildfire/index2.html

Fire Brochures for Public Distribution

http://www.interfacesouth.org/resources/brochure_all.html

Smoke Management Guide for Prescribed and 

Wildland Fire 

www.nwcg.gov/pms/pubs/SMG/SMG-72.pdf

Forest Encyclopedia Network

www.forestencyclopedia.net/

The Network connects scientific results and conclusions 

with management needs and issues. Focus areas include 

Southern Appalachian ecosystems, Environmental 

Threats, Southern Fire Science, and Southern Bioenergy.

Summary of the Restoration Goals for the National Forests 

in North Carolina

www.cs.unca.edu/nfsnc/restoration/priorities.htm

APPENDIX

Additional Fire Ecology Resources Available on the Internet
46

APPENDIX

Fire-related Publications for Hardwood, 

Upland Forests

Rainbow Series

Wildland Fire in Ecosystems: Effects of Fire on Fauna

www.fs.fed.us/rm/pubs/rmrs_gtr042_1.pdf

Wildland Fire in Ecosystems: Effects of Fire on Flora

www.fs.fed.us/rm/pubs/rmrs_gtr042_2.pdf

Wildland Fire in Ecosystems: Effects of Fire 

on Soils and Water

www.fs.fed.us/rm/pubs/rmrs_gtr042_4.pdf

Wildland Fire in Ecosystems: Effects of Fire on Air

www.fs.fed.us/rm/pubs/rmrs_gtr042_5.pdf

Wildland Fire in Ecosystems: Fire and 

Nonnative Invasive Plants

www.fs.fed.us/rm/pubs/rmrs_gtr042_6.pdf

Henri D. Grissino-Mayer’s Ultimate Tree-Ring 

Web Pages

web.utk.edu/%7Egrissino/my_page.htm

Fire and Fire Surrogates Study—Treatments 

for Ecological Restoration

www.fs.fed.us/ffs/index.html

Southern Forest Resource Assessment Chapter 25: 

Fire in Southern Forest Landscapes 

www.srs.fs.usda.gov/sustain/report/fire/fire.htm

Using Fire to Control Invasive Plants: What’s New, 

What Works in the Northeast

extension.unh.edu/Pubs/ForPubs/WPUFCI03.pdf

Fire Management Today “Fire Uses from the Past” 

Volume 64, No. 3

www.fs.fed.us/fire/fmt/fmt_pdfs/FMT64-3.pdf

Fire in the Southern Appalachians: Fuels, 

Stand Structure and Oaks

www.fs.fed.us/r8/boone/fire/fsa/index.shtml 

A Joint Fire Science Program project in Daniel Boone 

National Forest

Prescribed Fire and Oak Ecosystem Maintenance: 

A Primer for Land Managers

www.siu.edu/~forestry/faculty/groninger/Prescribed%20

Fire%20015.pdf

Proceedings from the 2000 Workshop on Fire, People, and 

the Central Hardwoods Landscape

www.treesearch.fs.fed.us/pubs/3762

Fire Learning Network Publications

tncfire.org/training_usfln_networkpubs.htm

Upland Oak Ecology Symposium: History, Current 

Conditions, and Sustainability

www.srs.fs.usda.gov/pubs/gtr/gtr_srs073/gtr_srs073.pdf

In particular, read the concluding paper, “Where Do We 

Go From Here?”

http://www.srs.fs.usda.gov/pubs/gtr/gtr_srs073/gtr_

srs073-blaney001.pdf

Dickinson, Matthew B., ed. 2006. Fire in eastern oak 

forests: delivering science to land managers. Proceedings 

of a conference; 2005 November 15-17; Columbus, OH. 

Gen. Tech. Rep. NRS-P-1. Newtown Square, PA: U.S. 

Department of Agriculture, Forest Service, Northern 

Research Station. 303 p.

www.treesearch.fs.fed.us/pubs/18405.

Hutchinson, Todd F., ed. 2009. Proceedings of the 3rd fire in 

eastern oak forests conference. Gen. Tech. Rep. NRS-P-46. 

Newtown Square, PA: U.S. Department of Agriculture 

Forest Service, Northern Research Station. 154 p. 

www.treesearch.fs.fed.us/pubs/17287.

— Beth Buchanan, Fire Ecologist

U.S. Department of Agriculture Forest Service, 

Southern Region
47

ADDITIONAL FIRE REFERENCES FOR THE SOUTHERN APPALACHIANS

Abrams, M.D. 2006. Ecological and ecophysiological attributes and 

responses to fire in eastern oak forests. In: Dickinson, M.B., ed. Fire 

in eastern oak forests: delivering science to land managers. Gen. Tech. 

Rep. NRS-P-1. Delaware, OH: U.S. Department of Agriculture Forest 

Service, Northern Research Station: 74-89. http://www.treesearch.

fs.fed.us/pubs/18434. [Date accessed: December 31, 2009].

Artman, V.L.; Sutherland, E.K.; Downhower, D.F. 2001. Prescribed 

burning to restore mixed-oak communities in southern Ohio: effects 

on breeding-bird populations. Conservation Biology. 15: 1423-1434.

Boerner, R.E.J.; Brinkman, J.A.; Sutherland, E.L. 2004. Effects of fire 

at two frequencies on nitrogen transformations and soil chemistry 

in a nitrogen-enriched forest landscape. Canadian Journal of Forest 

Research. 34: 609-618. http://www.treesearch.fs.fed.us/pubs/14588. 

[Date accessed: December 9, 2009].

Boerner, R.E.J.; Decker, K.L.M.; Sutherland, E.K. 2000. Prescribed 

burning effects on soil enzyme activity in a southern Ohio hardwood 

forest: a landscape-scale analysis. Soil Biology and Biochemistry. 32: 

899-908.

Brown, J.K. 2000. Introduction and fire regimes. In: Brown, J.K; 

Smith, J.K., eds. Wildland fire in ecosystems: effects of fire on flora. 

Gen. Tech. Rep. RMR- 42-vol. 2. Ogden, UT: U.S. Department of 

Agriculture Forest Service, Rocky Mountain Research Station: 1-8. 

http://www.treesearch.fs.fed.us/pubs/4554. [Date accessed: December 

26, 2009].

Clinton, B.D.; Vose, J.M.; Swank, W.T. 1993. Site preparation burning 

to improve Southern Appalachian pine-hardwood stands: vegetation 

composition and diversity of 13-year-old stands. Canadian Journal of 

Forest Research. 23: 2271-2277.

Clinton, B.D.; Vose, J.M.; Swank, W.T. 1996. Shifts in aboveground and 

forest floor carbon and nitrogen pools after felling and burning in the 

Southern Appalachians. Forest Science. 42: 431-441.

Clinton, B.D.; Vose, J.M.; Swank W.T. [and others]. 1998. Fuel 

consumption and fire characteristics during understory burning in a 

mixed white pine-hardwood stand in the Southern Appalachians. Res. 

Pap. SRS-12. Asheville, NC: U.S. Department of Agriculture Forest 

Service, Southern Research Station. 8 p.

Clinton, B.D.; Vose, J.M. 2000. Plant succession and community 

restoration following felling and burning in the Southern 

Appalachians. Tall Timbers Fire Ecology Conference Proceedings. 

21: 22-29.

Clinton, B.D.; Vose, J.M.; Knoepp, J.D.; Elliott, K.J. 2003. Stream nitrate 

response to different burning treatments in Southern Appalachian 

forests. In: Galley, K.E.M.; Klinger, R.C.; Sugihara, N.G., eds. 

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Rankin, W.T.; Herbert, N. 2014. Restoration in the Southern Appalachians: 

a dialogue among scientists, planners, and land managers. Gen. Tech. Rep. 

SRS-189. Asheville, NC: U.S. Department of Agriculture Forest Service, 

Southern Research Station. 48 p.

We address three key questions for restoration ecology in the Southern 

Appalachian Mountains. First, what is the role of fire, especially when 

used as a management tool for oak-dominated ecosystems? Second, what is 

the relationship between early successional habitat and biodiversity? And 

third, how do we regenerate oak ecosystems? To answer these questions, 

first, we examine the historic role of fire in the mountains, discuss its 

effects on forest resources, and summarize a strategy for restoring fire to 

ecosystems with a long history of fire exclusion. Second, we examine the 

relationship between early successional habitats and wildlife resources 

in the mountains, discuss the pattern and rate of natural disturbance, and 

provide suggestions for creating and maintaining early successional habitat. 

And third, we review current management for oak regeneration and discuss 

the implications for oak ecosystems in the absence of management. In 

addition to addressing current questions in restoration ecology, we provide 

an extensive bibliography of the scientific literature, especially for fire 

management. Our goal is to provide a concise and practical summary of 

the current restoration literature for use by forest planners and managers 

throughout the Southern Appalachian Mountains.

Keywords: Early successional habitat, natural disturbance patterns, 

natural fire, oak regeneration, prescribed fire, restoration ecology, Southern 

Appalachian Mountains, wildlife.
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