United States

Department of

Agriculture An Old-Growth Definition for

Forest Service Dry and Dry-Mesic Oak-Pine

Forests

Southern

Research Station

David L. White and F. Thomas Lloyd

A Section of the Old-Growth Definition Series
The Authors:

David L. White, Ecologist, Southern Research Station,

U.S. Department of Agriculture, Forest Service,

Clemson, SC 29634- 1003; and F. Thomas Lloyd, Research

Forester, Southern Research Station, U.S. Department

of Agriculture, Forest Service, Bent Creek, NC 28806.

Preface

Old growth is widely acknowledged today as an essential part of managed forests, particularly on public lands. However, this

concept is relatively new, evolving since the 1970’s when a grassroots movement in the Pacific Northwest began in earnest to

define old growth. In response to changes in public attitude, the U.S. Department of Agriculture, Forest Service, began

reevaluating its policy regarding old-growth forests in the 1980’s. Indeed, the ecological significance of old growth and its

contribution to biodiversity were apparent. It was also evident that definitions were needed to adequately assess and manage the

old-growthresource. However, definitions of old growth varied widely among scientists. To address this discrepancy and other

old-growth issues, the National Old-Growth Task Group was formed in 1988. At the recommendation of this committee, old

growth was officially recognized as a distinct resource by the Forest Service, greatly enhancing its status in forest management

planning. The committee devised “The Generic Definition and Description of Old-Growth Forests” to serve as a basis for

further work and to ensure uniformity among Forest Service Stations and Regions. Emphasis was placed on the quantification

of old-growth attributes.

At the urging of the Chief of the Forest Service, all Forest Service Stations and Regions began developing old-growth

definitions for specific forest types. Because the Southern and Eastern Regions share many forest communities (together they

encompass the entire Eastern United States), their efforts were combined, and a cooperative agreement was established with

The Nature Conservancy for technical support. The resulting project represents the first large-scale effort to define old growth

for all forests in the Eastern United States. This project helped bring the old-growth issue to public attention in the East.

Definitions will first be developed for broad forest types and based mainly on published information and so must be viewed

accordingly. Refinements will be made by the Forest Service as new information becomes available. This document represents

1 of 35 forest types for which old-growth definitions will be drafted.

In preparing individual old-growth definitions, authors followed National Old-Growth Task Group guidelines, which differ

from the standard General Technical Report format in two ways-the abstract (missing in this report) and the literature citations

(listed in Southern Journal of Applied Forestry style). Allowing for these deviations will ensure consistency across

organizational and geographic boundaries.

September 1998

Southern Research Station

P.O. Box 2680

Asheville, NC 28802
Table of Contents

Page

Introduction .......................................................... 1

Description of Forest-Type Group .................. 1

Piedmont ............................................................... 2

Distribution and composition ............................. 2

Old-growth forest exahples ............................... 3

Presettlement forests and disturbance ................ 3

Coastal Plain: Atlantic Section ........................... 5

Distribution and composition ............................. 5

Old-growth forest examples ............................... 6

Presettlement forests and disturbance ................ 6

Coastal Plain: Gulf Coast Section ...................... 6

Distribution and composition ............................. 6

Old-growth forest examples ............................... 6

Presettlement forests and disturbance ................ 7

Interior Highlands ............................................... 7

Distribution and composition ............................. 7

Old-growth- forest examples ............................... 7

Presettlement forests and disturbance ................ 8

Appalachian Mountains: Blue Ridge ................. 8

Distribution and composition ............................. 8

Old-growth forest examples ............................... 10

Presettlement forests and disturbance ................ 10

Appalachian Mountains: Plateau ....................... 1 1

Distribution and composition ............................. 1 1

Old-growth forest examples ............................... 1 2

Presettlement forests and disturbance ................ 12

Appalachian Mountains: Valley and Ridge ...... 13

Distribution and composition ............................. 13

Old-growth forest examples ............................... 13

Presettlement forests and disturbance ................ 13

Forest Composition: Synthesis across Regions.. 14

Page

Narrative of Old-Growth Conditions ............... 14

Living Tree Component ........................................ 23

Size structure ....................................................... 23

Age structure ....................................................... 23

Canopy structure .................................................. 24

Dead Tree Component: Coarse Woody Debris . . 26

Forest Dynamics and Ecosystem Function ..... 27

Presettlement Disturbance Regimes .................... 27

Disturbance continuum ........................................ 27

Fire frequency ..................................................... 27

Patch size .............................................................27

Ecological Processes .............................................. 28

Postsettlement changes ........................................ 28

When Does Old Growth Begin? ........................... 29

Information Gaps ........................... ...................... 29

Representative Old-Growth Forests ................. 30

Acknowledgment ................................................... 30

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

Appendix A ............................................................. 35

Appendix B ............................................................. 42

An Old-Growth Definition for Dry

and Dry-Mesic Oak-Pine Forests

David L. White and F. Thomas Lloyd

Introduction

Dry and dry-mesic oak-pine forests are widely distributed

throughout the Eastern and Southern United States from

New Jersey to Texas. Shortleaf pine is the predominant pine

species of this forest type in the Piedmont, Gulf Coastal

Plain, and Interior Highlands, while pitch pine is the

predominant pine species in the Appalachians. White oak is

the predominant oak species across the range of old-growth

oak-pine sites, although scarlet, chestnut, and other

hardwood species share dominance on some sites.

Oak and pine species associated with this type have been

prevalent in the Eastern United States for thousands of years

(Delcourt and Delcourt 1987). The oak-pine type has been

described as ephemeral on a given site and is maintained in

a changing landscape mosaic by fire and other disturbance

(Buckner 1989). Forests of this type do not adhere to the

steady state or equilibrium concept of forest dynamics

historically associated with “virgin” or old-growth forests. In

addition, in defining old growth, we acknowledge that

multiple ecological outcomes are possible for comparable

sites (Oliver 1981, Sprugel 1991). The type, intensity, and

frequency of disturbance and its interaction with the biotic

and abiotic components of a specific site determine the

“outcome.” A greater understanding of these interactions

should clarify management options for recognizing,

creating, and maintaining old-growth types on the landscape.

Due to extensive postsettlement land clearing and soil

disturbance in the Eastern United States, examples of oldgrowth forests are rare. To characterize the dry and drymesic oak pine type, we used recent studies and historical

accounts of composition. In addition, we compiled

information from existing descriptions of representative oldgrowth stands of this type. Across these old-growth sites,

information was sparse. Therefore, one site in the South

Carolina Piedmont was sampled intensively to obtain one

full set of old-growth attribute data (White and Lloyd 1995).

Additional data from this site will be presented here for the

fust time.

In this paper we describe the dry and dry-mesic oak-pine

old-growth type for each of six physiographic provinces

with an emphasis on composition, distribution (including

presettlement), old-growth examples, and disturbance

regimes. Specific attributes from a range of old-growth sites

are presented in a standardized table and discussed; then,

presettlement disturbance regimes and ecological processes

associated with this type are discussed. Finally, we address

the following questions regarding this oak-pine forest type:

(1) when is a forest considered old growth, and (2) what is

the minimum area needed to maintain old-growth forests?

Description of Forest-Type Group

The starting point for defining the dry and dry-mesic oakpine type relates to soil moisture and the relative

composition of oaks and pines. The soil moisture class for

this type is defined as dry and dry-mesic, corresponding to

soils that are somewhat excessively drained to well drained.

Regarding the relative composition of oak and pine, we

currently define the upper limit of pine composition to be

~50 percent. Our reason for this is because the original oldgrowth classification for the Eastern United States (35

types) does not include a “pine-oak” forest type (pine

composition >50 percent) that occurs on dry-mesic sites. To

exclude a pine-oak type presumes that pine-dominated forest

types were not represented on dry-mesic sites within the

Eastern United States under presettlement disturbance. We

do not consider this a sound assumption, especially

considering that pine dominance on these sites has been

described or implied for the southern Piedmont (Nelson

1957, Phunmer 1975), upper section of the Atlantic Coastal

Plain (Sargent 1884), portions of the Gulf Coastal Plain

(Mohr 1901, Harper 1943, Delcourt 1976, Schafale and

Harcombe 1983, Martin and Smith 1991), and portions of

the Ozarks and Ouachitas (Foti and Glenn 199 1, Tucker

1991). Defting an upper limit for pine composition (or a

lower limit for oak) is somewhat arbitrary. From the Society

of American Foresters (SAF) forest cover-type descriptions

(Eyre 1980), “A species must comprise at least 20 percent of

the total basal area to be used in the type name.” Interpreting

“species” broadly as either pines or oaks, we define the

composition of the dry and dry-mesic oak-pine type as

containing at least 20 percent pine basal area and at least 20

percent oak basal area. Using this designation, the upper

limit for pine composition is 79 percent.

Dry and dry-mesic oak-pine forests constitute a large part of

the eastern deciduous forest, extending from southern
Missouri to east Texas in the West across to the Atlantic

coast from southern New York to northern Florida. This

type occurs in the following physiographic provinces:

Coastal Plain, Piedmont, Blue Ridge, Valley and Ridge,

Appalachian Plateau, Ozark Plateau, and the Ouachitas.

Most forests occur on coarse textured soils on ridges and

south-facing slopes in the mountains and droughty uplands

of the Piedmont and Coastal Plain. The inherent dryness of

these sites makes them particularly prone to fire.

Principal species’ vary widely depending on latitude,

topography, geography, and disturbance. On dry sites,

forests can include shortleaf pine, Virginia pine, Table

Mountain pine, pitch pine, post oak, chestnut oak, scarlet

oak, and blackjack oak. On dry-mesic sites, loblolly pine,

white oak, southern red oak, black oak, mockemut hickory,

pignut hickory, red maple, and blackgum are more

prominent. Since most of the above species are restricted to

certain physiographic regions, not all of these species

necessarily co-occur.

Dry and dry-mesic oak-pine forests are similar to the

following SAF cover types: shortleaf pine (75), shortleaf

pine-oak (76), Virginia pine-oak (78), Virginia pine (79),

loblolly pine-shortleaf pine (80), loblolly pine (8 I), and

loblolly pinezhardwood (82).

The composition of the dry and dry-mesic oak-pine type

(type 25) is similar to that of two other old-growth forest

types-xeric pine and pine-oak forests and woodlands (type

24) and dry-mesic oak forests (type 2 1). Types 24 and 25

may occur adjacent to each other along a moisture gradient,

with type 24 occupying more xeric sites along ridges. Types

2 1 and 25 may develop on some of the same site types

depending on the type and intensity of disturbance.

Since few old-growth examples of dry and dry-mesic oakpine forests exist, both recent studies and historical accounts

of composition were used to describe this forest type.

Recent studies consist of descriptions of “natural” vegetation

within a region. Extensive postsettlement land clearing and

soil disturbance in the East make it necessary to use

historical documents and original land survey records to

reconstruct pre- or early-settlement composition even

though the reliability of these approaches is somewhat

variable.* It is also important to acknowledge the dynamic

’ See appendix A for a list of scientific and common plant names as well as

their presence in old-growth and non-old-growth forest stands. Scientific

names are given in the text only if they were not included in appendix A or

if used in reference to a number of species within one genus as in

Vaccinium species. Nomenclature is from Kartesz and Kartesz (1980).

’ Frost, C.C. 1997. Presettlement vegetation and natural tire regimes of the

Savannah River Site. 179 p. Unpublished report. On file with: Savannah

River Natural Resource Management and Research Institute, USDA Forest

Service, New Ellenton, SC 29809.

nature of presettlement forests, which makes their

assessment somewhat arbitrary. Furthermore, descriptions of

forests at the time of settlement may describe forests

significantly impacted by Native American population

decline (i.e., less fire) as related to exposure to diseases

introduced by explorers 200 years before settlement

(DeVivo 199 1).

Given the broad distribution of the dry and dry-mesic oakpine type, we have chosen to describe this forest type by

physiographic regions. For each physiographic region given

below, we describe the old-growth type in the context of

(1) distribution and composition, (2) old-growth forest

examples (composition and location), and (3) presettlement

forests and disturbance.

Piedmont

Distribution and composition-The Piedmont extends

from the palisades of the Hudson River to Alabama. Few

descriptions of vegetation composition exist for the entire

Piedmont region. Although Godfrey’s (1980) work was

purely descriptive and did not consider pine as a mature

forest species, his description of initial woody succession

provided insight into the distribution of yellow pine species

across the region. Pines share dominance with hardwoods

during early succession from Maryland to northern South

Carolina but dominate from South Carolina southward.

Virginia pine is predominant in the northern Piedmont,

sharing dominance with shortleaf pine in Virginia. Virginia

pine decreases in importance from Virginia southward,

occupying only the upper Piedmont in South Carolina.

Shortleaf pine is dominant from North Carolina southward,

with loblolly pine increasing in importance from the fall line

westward where it becomes dominant in the Alabama

Piedmont.

Braun (1950) considered most of the Piedmont a part of the

oak-pine region with the exception of monadnocks, which

were considered “Blue Ridge outhers,” and more

vegetationally similar to oak-chestnut forests. Within the

Virginia Piedmont, a shift from the oak-pine to the oakchestnut region is related to a shift from red and yellow soils

(Cecil Series) to gray-brown forest soils (Chester Series)

(Braun 1950). The oak-chestnut section in southern Virginia

starts as a narrow, hilly belt adjacent to the Blue Ridge and

contains an abundance of monadnocks (Fenneman 1938).

North of the Potomac River is the Piedmont Lowland. Here,

American chestnut, oaks, and hickories once dominated this

area as well as the narrow, hilly belt, with pitch and Virginia

pines as occasional associates. The intensity of

postsettlement disturbance precludes a clear picture of early

vegetation in the northern Piedmont.
In the southeastern Piedmont, we refer to site or vegetation

studies, or both, l?om North Carolina3 and South Carolina

(Jones 1988) to provide insight into the distribution of the

dry and dry-mesic oak-pine type. Peet compiled a tabulation

of synonymous communities for the North Carolina

Piedmont from various sources. This tabulation and the

original studies allow us to infer which oak communities

occupied sites where the dry and dry-mesic oak-pine type

could develop, given sufficient disturbance (e.g., fire, wind

and ice storms, and insects). The following tabulation shows

similar community types that occupy sites where old-growth

oak-pine may occur:

Community synonymy for North Carolina Piedmont

Peet and Schafale and

Christensen 1980 Oostiug 1942 Weakley 1990

Dry mesotrophic

Clayey phase

Sandy phase

Dry oligotrophic

Dry eutrophic

White oakblack oakred oak

White oak

White oakpost oak

Dry-mesic oakhickory

Dry oak-hickory

Basic oak-hickory

In South Carolina, Jones (1988) developed a predictive

landscape ecosystem classification that integrates landform,

soil, and late successional vegetation. While Jones’ system

predicts the late successional hardwood vegetation type

from soil and landform variables, it is important to

acknowledge that multiple ecological outcomes are possible

on a given site unit. Different forest types will develop for

each combination of soil type, landform, and disturbance

regime. Based on Jones’ descriptions, literature information,

and our sampling, old-growth dry and dry-mesic oak-pine

could develop on xeric, subxeric, and intermediate sites with

the center of distribution on subxeric sites (table 1).

Old-growth forest examples-Few old-growth oak-pine

stands have been documented in the Piedmont. Oosting

(1942) described a relatively undisturbed white oak-post oak

type in the Duke Forest of North Carolina that contained

shortleaf pine (appendices A and B). He considered this

type a variant of the oak-hickory climax. Shortleafpine was

the next most important species (basal area) in stands

dominated by white and post oaks, followed by hickory

species, scarlet and northern red oaks with dogwood

prominent in the understory. Minor associates included

3 Peet, Robert K. 1994. Community synonymy tables and data. 5 p.

Unpublished report. On file with: University of North Carolina, Chapel

Hill. NC 275 14.

black and blackjack oaks, sourwood, blackgum, and red

maple. Shrubs included deerbeny, black-haw, rusty blackhaw, other Viburnum and Vaccinium species, wild azalea,

strawberry bush, and fringe tree. Vines included muscadine,

greenbrier, trumpet honeysuckle, and Virginia creeper.

Herbs included broom-sedge, Panicum species, Carex

species, wild ginger, five-finger, hawkweed, pipsissewa,

bedstraw, and dwarf Dutchman’s-pipe.

In South Carolina, we sampled the John de la Howe (JDLH)

tract-a relatively undisturbed 200-year-old oak-pine stand

on subxeric and intermediate sites where insects had

sporadically killed pines (white and Lloyd 1995,

appendices A and B). Shortleaf and loblolly pines and white

oak were the dominant species followed by various hickory

species, yellow-poplar, post oak, southern red oak, red

maple, sourwood, black oak, and elm. Dogwood was the

most important understory species. Others included eastern

hophornbeam and Florida or southern sugar maple. Vine,

shrub, and herb species given in order of decreasing

frequency included muscadine, Smilax species, partridgeberry, bedstraw, wild ginger, strawberry bush, Virginia

creeper, Carex species, miscellaneous grasses, perfoliate

bellwort, dwarf Dutchman’s-pipe, pipsissewa, Viburnum

species, Solomon’s-seal, deerberry, wild yam, sparkleberry,

and others. Red maple was the most abundant tree seedling,

followed by eastern hophombeam, dogwood, pines

(shortleaf and loblolly), black cherry, white oak, chalk

maple (Acer leucoderme Small.), pignut hickory, black oak,

and blackgum. The pine component in these stands is

declining, and, even though seedling abundance was

moderately high, no pines were represented in the 0.8- to

3.9~inch [ 1.8- 9.9 centimeter (cm)] diameter at breast height

(d.b.h.) class, suggesting successful recruitment is unlikely.

Lack of fire in this stand has likely resulted in the

predominance of mesic understory species such as red and

southern sugar maple, eastern hophornbeam, and others

(Orwig and Abrams 1994).

Presettlement forests and disturbance--The composition

of southeastern Piedmont forests before European settlement

has been described (Nelson 1957, Brender 1974, Plummer

1975, Skeen et al. 1980, Peet and Christensen 1987, Skeen

et al. 1993, Barden 1997), but no clear agreement exists.

From these papers and several historical accounts of the

Georgia, North Carolina, and South Carolina Piedmont

[Drayton 1784, Mills 1826, Ruffm 1843, Logan 1859,

Hammond 1883, Schoepf 1911, Bartram 1958, Lawson

1967, Clark 1973 (as cited in Michaux 1802)], the landscape

consisted of relatively open, hardwood- dominated forests

with scattered pines, extensive prairies and savannahs, and

large stretches of cane brakes along streams and lowlands.

Burning by Native Americans was credited for the

3
Table l-Late successional community type, soil, and landform characteristics from the South Carolina Piedmont

Landform Ecosystem Classification (Jones 1988) corresponding to potential sites for old-growth dry and dry-mesic

oak pine

Site unit and community type Distinguishing soil characteristics Landform description

Xeric

Post oak, black oak,

and lowbush blueberry”

Subxeric

White oak-scarlet oakdeerberty

Heavy clay subsurface horizon 12 in.

(3 1 cm) from surface or bedrock

within 24 to 36 in. (61 to 91 cm)

Clay or sandy clay subsurface horizons

at 12 to24 in. (31 to 61 cm)

Or:

Exposed ridge flats, upper

slopes of any aspect

Ridge flats, upper slopes of

any aspect or midslope with

a southern aspect

Heavy clay horizons within 12 in.

(3 1 cm) of surface or bedrock

within 24 to 36 in. (61 to 91 cm)

Less exposed midslopes

with north or east aspects

Intermediate

White oak-northern red oakfalse Solomon’s- seal

Sandy clay loam or clay loam at 12

to 24 in. (61 to 91 cm)

Midslopes with north or east

aspects or mid to lower slopes

with southern aspects

a Vaccinium vacillans TOIT.; other species are given in appendix A.

maintenance of this early landscape (Rostlund 1957, Barden

1997). A case could be made, at least for parts of the South

Carolina Piedmont, that pines became more abundant in the

Piedmont after the mid- 1700’s by establishing on the more

open areas after the Cherokees sold much of their land to the

British (compare mid- 1700’s accounts by Logan 1859 and

1980 in reference to pine by Drayton 1784, Mills 1826, and

Ruffm 1843). Pine establishment on abandoned agricultural

land started in the late 1700’s.

The Piedmont, as described by Peet and Christensen (1987),

was dominated by oaks, hickories, and other hardwoods

with loblolly and shortleaf pines found only occasionally,

usually on exceptionally poor sites or where some

disturbance (windfall, fire, or aboriginal cultivation) had

created suitable conditions for pine. Skeen et al. (1980)

suggested that oak-hickory-yellow-poplar was the “typal”

forest climax in the southeastern Piedmont, with oak-hickory

and oak-hickory-pine as edaphic and moisture-controlled

variants of this type. Brender (1974) describes the “original”

forest in the lower Piedmont as the “oak-hickory climax

type” with other associated hardwoods and a “scattering of

shortleaf and loblolly pines. ” The presumption that pine was

only found occasionally in the Piedmont contrasts with the

descriptions for the Georgia and Alabama Piedmont.

Nelson (1957) described the “original” forest of the Georgia

Piedmont as ranging from pure pine to pure hardwoods.

Oak-hickory types were more abundant in the “red lands”

(35 to 40 percent of the Georgia Piedmont), whereas

shortleaf pine and post, white, and red oaks occupied the

“gray sandy lands” (45 percent of the region). These latter

soils are generally described as gray sand and gravelly and

probably are the uneroded Cecil, Appling, and Durham

soils. Pine dominated the remaining 15 percent of the

Piedmont, which is the Elberton granitic formation. Nelson

indicated that pines were more prevalent west of the

Atlantic-Gulf divide and generally decreased from the

sandhills to the mountains. Plummer (1975) used survey

records of the Georgia Piedmont and Blue Ridge from 1733

to 1832 to provide more specific information. Pine was

especially abundant on steep slopes and upland soils of

micaceous origin. Soils with higher clay content supported

more post oak. The ratio of oak:pine:hickory was 5312318 on

over one-half million acres surveyed.

Pine species were also a significant component of the

“original” forests of the Alabama Piedmont (Mohr 190 1,

Harper 1943), with shortleaf occurring in “dry woods” along

with southern red oak, post oak, hickory species, and black

oak. Dogwood, black-haw, and hawthome were common

understory species. Longleaf pine and loblolly pine were

also significant components of this region in Alabama. Mohr

(1901) also described a mountainous area where longleaf

pine dominated in open pine woodland along rocky ridges

mostly below 2,000 feet [610 meters (m)]. Blackjack oak

was an important associate, being replaced by chestnut oak,
American chestnut, and pignut hickory. On the deeper soils

of slopes and ridges, hardwoods predominated with

shortleafpine as an occasional associate. This mountainous

section, including Rebecca and Talladega Mountains,

consists of both slate and quartzite rocks. Due to its location

and parent material, there is disagreement as to which

physiographic province this section belongs-Valley and

Ridge, Blue Ridge, or Piedmont 4 (Femreman 1938,

Hodgkins et al. 1979). Braun (1950) described this section

as an area where American chestnut, chestnut oak, and

longleaf pine intergrade.

Whereas fire or other disturbances likely maintained pine as

a component of forests in the Piedmont, we understand little

of the spatial and temporal aspects of presettlement

disturbance (human and nonhuman) in the Piedmont. Such

understanding would provide insight into the relative

abundance of the oak-pine type in this region. Frost (1995)

estimated the presettlement fire frequency to be 4 to 6 years.

He suggested that the influence of Native American fire was

greater in the Piedmont than in the Coastal Plain (where

lightning ignitions predominate) because of differences in

fire compartment size.’

Differences in fuel moisture between the two regions also

contribute to variations in fire occurrence. Fuel moisture is

generally higher in the Piedmont, largely due to clay soils

that retain water near the surface (Christensen 198 1). It

seems safe to deduce that the oak-pine type has been a

component of the Piedmont for millennia. More research is

needed, however, to characterize the extent of its

presettlement distribution.

Coastal Plain: Atlantic Section

Distribution and composition-The Atlantic Coastal Plain

extends from Florida to New Jersey. Specific information on

the dry and dry-mesic oak-pine type’s occurrence in this

section is sparse. Most of the Coastal Plain north of the

James River in Virginia was considered by Braun (1950) to

be part of the oak-pine forest region, whereas that area south

of the James River was considered southeastern evergreen

forest. The upland forests of the eastern and western shores

of Maryland resembled those of the Piedmont further south,

according to Braun, with loblolly pine in association with

Virginia pine or deciduous species.

4 Personal communication. 1994. Henry McNab, Research Forester,

Southern Research Station, USDA Forest Service, Bent Creek Research

and Demonstration Forest, 1577 Brevaxd Road, Asheville, NC 28806.

5 Frost (1995) defines “fire compartment” as a unit of the landscape with

no natural firebreaks, such that a lightning ignition in one part would be

likely to burn the whole unit unless there were a change in weather or fuel

conditions.

On the Coastal Plain of New Jersey, pine-oak and oak-pine

types are prevalent and include pitch and shortleafpines,

several oaks, and understory ericaceous species. The dry and

dry-mesic oak-pine type corresponds to the pine-oak type

described by Lutz (1934). This type is somewhat similar to

the pitch pine forests of Southern Appalachian ridges

(Whittaker 1979), but it also has species common to the

southern Atlantic Coastal Plain (Christensen 1988a). Breden

(1989) describes a dry oak-pine forest of the outer Coastal

Plain of New Jersey as a closed canopy community where

dominant tree heights range from 36 to 50 feet (11 to 15 m)

to 75 to 102 feet (23 to 3 1 m) depending on fire history. The

dominant oak species are black, chestnut, scarlet, post, and

white, with pitch and shortleaf pines as important associates.

Huckleberry, dangleberry, bear oak (Quercus iliczjbliu

Wang.), and lowbush blueberry occur in some of these

stands. Breden further noted a scarlet oak-shortleaf pine

subtype of the dry oak-pine forest as occurring on the

eastern half of the Pine Barrens where scarlet and black oaks

and shortleaf pine dominate. Little and Moore (1949)

described the New Jersey pine region as consisting of

“subclimax” stands of hardwoods and pine that have

resulted from heavy cuttings and severe fires, the latter

occurring at intervals of 15 to 40 years.

The dry and dry-mesic oak-pine type is uncommon on the

southeastern Coastal Plain but fire suppression, the degree

of postsettlement disturbance, and the scarcity of literature

information precludes conclusive determination on this

subject. Barry (1980) refers to scrubby oak barrens

occupying the most xeric sites in the South Carolina

sandhills and xeric pine-hardwoods in somewhat more mesic

areas where sands are not as deep. In the latter, loblolly pine

occurs with post, southern red, dwarfpost, black, water (Q.

nigra L.), and blackjack oaks and pignut and mockemut

hickories. Barry describes similar communities for other

sections of the Coastal Plain, but their occurrence appears

infrequent.

Vegetation descriptions from Georgia (Pehl and Brim

1985), South Carolina (Barry 1980, Nelson 1986), and

North Carolina (Schafale and Weakley 1990) are not clear

on the occurrence of this type in the middle and lower

Coastal Plain sections. In a synthesis of vegetation studies of

the southeastern Coastal Plain by Christensen (1988a), there

is no mention of a type corresponding to dry and dry-mesic

oak-pine, except for a statement that mid-Atlantic flatwoods

are roughly similar to the upper slope pine-oak forests

described by Marks and Harcombe (198 1) for east Texas

(see Coastal Plain: Gulf Coast Section). The latter is

considered to be similar to the dry and dry-mesic oak-pine

type.

5
Old-growth forest examples-No documented examples

of old-growth oak-pine exist to our knowledge.

Presettlement forests and disturbance--Historical

references to this type are limited. In the Coastal Plain of

southern Delaware, pine was considered an insignificant

component of the “original” forest (Sterrett 1908). In the

southeastern Coastal Plain, Bartram (1958) lists shortleaf

and loblolly pines occurring with various Coastal Plain and

Piedmont oaks approximately 50 miles [80 kilometers (km)]

inland from Savannah, GA, but shortleaf is rarely

encountered in the middle and lower Coastal Plain today.

The most definitive early reference is from Sargent (1884),

who described a transition area corresponding to the

sandhills and upper Coastal Plain region as “shortleaved and

loblolly pine intermixed with hardwoods in about equal

proportion and scattered long leaved pine.” Frost (1993)

views these transitional types as rare remnants of savanna

and woodland fire types with a species-rich herb layer, in

contrast to the postlogging, fire-suppressed, successional,

mixed pine-hardwood forests prevalent today. From Frost’s

(see footnote 2) descriptions of presettlement forests of the

South Carolina sandhills, the dry and dry-mesic oak-pine

type is likely best represented in the fire tension zones

between upland longleaf and nonpyrophytic woodland

below. Although sometimes relatively narrow in the

sandhills, these fire tension zones can be much broader in

other parts of the Coastal Plain (see footnote 2). These

pyrophytic woodlands include shortleaf pine as a key

species along with longleaf, white and post oaks, and,

occasionally, loblolly. Shortleafpine is rarely mentioned in

more recent descriptions of these areas.

Sargent’s (1884) description of a transition area in the upper

Coastal Plain and sandhills is the most significant evidence

for the occurrence of the dry and dry-mesic oak-pine type in

the region. The disturbance dynamics that maintained this

type on the landscape might be more similar to that

prevailing in certain parts of the Gulf Coastal Plain, which

will be discussed in more detail later. The extent of the dry

and dry-mesic oak-pine type’s distribution in the mid- and

lower Coastal Plain sections is unknown.

Coastal Plain: Gulf Coast Section

Distribution and composition-The dry and dry-mesic

oak-pine type is a more significant component of the Gulf

Coastal Plain than the Atlantic Coastal Plain, occurring from

Florida to Texas. The belted character of this region in the

form of inner lowlands and cuestas and other low-ridge

landforms (Bowman 19 11, Fenneman 1938), the associated

diversity of soil types, and differences in settlement history

account for much of the difference in this type’s distribution

in the two Coastal Plain regions. Cuestas and other hills

create strong environmental gradients which, coupled with

soil characteristics, result in highly pyrogenic longleaf

stands on sandy or sandy loam ridges, shortleaf pinehardwood stands on upper to midslopes on clay soils,

whereas loblolly pine-hardwood occupy lowland silt loam

soils (Mohr 190 1, Martin and Smith 199 1). Wide variation

in composition across this gradient is related to fire

frequency and intensity (Martin and Smith 199 1). Primarily,

the shortleaf-hardwood type represents the dry and drymesic oak-pine type. Unlike most of the Atlantic Coastal

Plain, shortleaf pine is an important species across much of

the Gulf Coastal Plain, both presently and historically (Mohr

1901; Harper 1920,1943; Delcourt 1976; Martin and Smith

1991).

Marks and Harcombe (198 1) described composition in the

Big Thicket of Texas. Stands on moderate to well-drained

sandy loams on upper and midslopes of low ridges were

described as dry to dry-mesic. The upper slope pine-oak

forests were dominated by shortleaf, loblolly, southern red

oak with longleaf, post oak, blackjack oak, and sweetgum

(Liquidambar styracifua L.) as associates. Midslope oakpine forests were dominated by loblolly pine, white oak,

southern red oak, shortleaf pine, with equal representation of

red maple, sweetgum, blackgum, and American holly (Z/ex

opaca Ait.). Sweetgum was also present on upper slopes.

Shortleaf and loblolly were more prevalent on the upper and

midslope, respectively. Ridgetops of deep sands were

dominated by longleaf pine-scrub oak. The authors note the

similarity of the upper and midslope communities described

above to previous descriptions of oak-pine and pinehardwood types of the upper Coastal Plain and Piedmont

(Braun 1950, Nelson 1957, Kuchler 1964, Delcourt 1976,

Golden 1979). A similar moisture gradient with associated

plant communities has been described for the Louisiana

Coastal Plain (Martin and Smith 199 1).

Old-growth forest examples-In the belted section of the

Louisiana Coastal Plain on the Kisatchie National Forest,

Martin and Smith (199 1) described two old-growth shortleaf

pine-oak-hickory stands (appendices A and B). Shortleaf

pine; post, southern red, and white oaks; loblolly pine; and

mockemut hickory dominated these stands. The midstory

dominants were eastern hophornbeam and dogwood with

red maple, blackgum, and hawthome. Understory shrubs

include yaupon, horse-sugar, and wild azalea. Blueberry was

the dominant shrub in one of the stands. The most abundant

herbs were partridge-berry, Carex species, Panicum species,

Uniola species with lesser amounts of goldenrod,

Desmodium species, Aster species, elephant’s-foot, and little

bluestem. Martin and Smith also described the composition

of the “shortleaf pine-oak-hickory natural community type”

based on historical and current information, which included

species not found in the old-growth stands (see appendix A).

They noted the shortleaf pine-oak understory was not as

species rich as the adjacent longleaf pine community.

6
Presettlement forests and disturbance--Descriptions of

“original” or early settlement forests (before extensive

European disturbance) for Alabama (Mohr 190 1, Harper

1943) and Mississippi (Hilgard 1860) report the widespread

occurrence of shortleaf pine-mixed oak throughout the

middle and upper Coastal Plain. This community was

commonly found on dry uplands with sandy clay to sandy

soils and prevalent in the upper or hilly Coastal Plain, post

oak flatwoods, red hills, and upper portion of the longleaf

pine belt of the middle Coastal Plain (pine hills). Shortleaf

was associated with southern red and post oaks and loblolly

pine, along with dogwood in the understory. Scarlet oak,

blackjack oak, mockemut hickory, and white and black oaks

were also common. Longleafpine communities, occurring

on dry, sandy ridges, were commonly adjacent to shortleaf

pine-oak communities in this region.

Hilgard (1860) described shortleaf pine and associated

hardwood species as being significant components of the

northeast pine hills, flatwoods, yellow loam region (includes

red hills), central prairie, and upper section of the longleaf

pine region (pine hills) in Mississippi. Longleafpine was

largely confined to the pine hills region of the middle

Coastal Plain, which contrasts with its more widespread

distribution in Alabama. Hilgard mentions American

chestnut as an-occasional associate of the oak-shortleafpine

type in addition to the associated species mentioned above

for Alabama. Shortleaf pine and associated hardwoods were

generally found on sandy ridges or uplands between

drainages. In areas with less topographic relief or with more

clayey soils, hardwoods increased in importance.

Braun (1950) described the oak-pine region of the Coastal

Plain of Texas, Louisiana, and Arkansas as rolling to hilly

with shortleaf as the dominant pine in the northern section

“originally” and loblolly as the dominant in the southern

section where longleaf is also prevalent. Associated oaks

were post, blackjack, and southern red. Braun also described

a transition area between the oak-pine and oak-hickory

forest regions in southern Arkansas where pine is a

component of oak-hickory forests that contain white,

chestnut, southern red, and Shumard oaks (Q. shumardii

Buckl.), as well as pignut, mockemut, and bittemut hickory

[ Carya cordzjixmis (Wangenh.) K. Koch].

Studies of presettlement forest composition in northern

Louisiana (Delcourt 1976) and east Texas (Schafale and

Harcombe 1983) suggest where longleaf, shortleaf, and

loblolly pines likely occurred on the landscape. Shortleaf

pine probably dominated the northern part of the hilly

Coastal Plain section north of the Red River in Louisiana

whereas longleaf dominated the southern part. The

abundance of pine in this region was attributed to natural

fires and Indian activities (Delcourt 1976). In the

presettlement landscape of the Big Thicket area of east

Texas, prairies were more extensive than they are today, and

pine species were prevalent in most community types,

including mesic hardwood forests (Schafale and Harcombe

1983). An upland pine type (95 percent pine) was likely

longleaf dominated, whereas the intermediate pinehardwood (57 percent pine) and white oak-red oak-pin oak

(Q. palustris Muenchh.) types (32 percent pine) probably

contained loblolly and shortleaf. The authors acknowledged

fire and other disturbances as favoring pine abundance in the

more mesic forests, but the scale of disturbance was not

discussed. Since the Big Thicket experiences many tropical

storms, it is conceivable that the 1800’s presettlement pine

composition was high because of storm-related damage

followed by intense fires due to heavy fuel loads.

Frequency of lightning fire was estimated to be 5 to 15 years

in shortleaf pine-oak forests in Louisiana and 5 years or less

in adjacent longleafpine communities (Martin and Smith

199 1). Canopy level disturbances (blowdowns) in the

shortleaf-oak stands were likely small to moderate in size

based on the uneven-aged structure of these stands. The

authors estimated that over one-half of the “original”

shortleaf pine-oak-hickory forest was now in mixed

hardwood-loblolly, whereas approximately one-third of the

“original” longleaf pine forest was now in shortleaf-oakhickory. Marks and Harcombe’s (198 1) study describing the

Big Thicket of Texas supports Martin and Smith’s concept

that the pyrogenic longleaf pine community functions as a

fire source for adjacent, somewhat less flammable,

communities. With more frequent fne, longleaf pine

increases on upper slopes, whereas shortleafpine increases

on midslopes with a decline in mesic hardwood species.

With less fire, loblolly and other mesic species move or

“creep” upslope, a phenomenon observed in the Piedmont

and other regions where loblolly is present.

Interior Highlands

Distribution and composition-This region includes the

Ouachita Mountains and the Ozark Plateau that Braun

(1950) characterized as the oak-hickory forest region. The

Ozark Plateau consists of a dissected plateau and the higher,

more rugged Boston Mountains on the southern end.

Limestones, cherts, and dolomites are the predominant

parent materials of the plateau, whereas sandstones and

shale are common in the Boston Mountains. The Ouachitas

consist of shales, sandstones, quartzites, and cherts.

Although Braun viewed oak-hickory as the prevailing type,

pine and pine-oak communities occurred on southern slopes

and ridges. Shortleaf was more abundant in the Ouachitas

than in the Ozark Plateau. Species associated with shortleaf

include post, blackjack, white, black, and southern red oaks

and hickory species on upland sites.

Old-growth forest examples-Several old-growth stands

have been characterized for the Interior Highlands (Dale and

Watts 1980, Fountain and Sweeney 1985, Fountain 1991)

7
(appendices A and B). Most of the Ouachita Mountains are

oriented east-west, resulting in distinct vegetation types on

the north- and south-facing slopes. In the Roaring Branch

Research Natural Area (Roaring Branch RNA), north- and

lower south-facing slopes were dominated by white oak,

northern red oak, mockemut hickory, and other hardwoods

(Fountain and Sweeney 1985). South slopes, at mid and

upper slope positions, were dominated by shortleafpine,

oaks (northern red, white, and black), and mockemut

hickory with shortleaf more abundant on midslopes.

Blackjack oak and black hickory were associates on southfacing ridges. Understory species included lowbush

blueberry, sparkleberry, and eastern hophombeam on mid

and upper slopes, while Smilax species, Virginia creeper,

and sassafras were found on upper south-facing slopes only.

In the Hot Springs National Park (Hot Springs NP), the dry

and dry-mesic oak-pine type is best exemplified by the oakhickory-pine type (mesic and xeric subtypes) as described

by Dale and Watts (1980). The mesic subtype occurs on

lower north-facing slopes and flats where shortleaf and

white oak share dominance. Other species include

mockemut hickory, blackgum, and southern red, post, and

black oaks. The xeric subtype occupies south and southeast

slopes and occasionally steep, northern exposures and is

dominated by shortleaf followed by post and blackjack oaks,

black and mockemut hickory, and white oak.

The Lake Winona Research Natural Area (Lake Winona

RNA) is an example of an uneven-aged, old-growth,

shortleaf pine stand in the Ouachita Mountains (Fountain

and Sweeney 1987, Fountain 1991). Unlike most of the

Ouachitas, ridges in this area are north-south oriented.

Vegetation was similar on east and west ridges, with

shortleaf dominant. Uplands consisted of mostly clayey soils

over shale where white oak was the dominant hardwood.

Hardwood composition is similar to that reported for Hot

Springs NP and Roaring Branch RNA. Lowbush blueberry

was the most important shrub species.

Presettlement forests and disturbance-The Ouachita

Mountain landscape at the time of settlement contained an

abundance of shortleaf pine, which tended to dominate on

sandstone substrates, whereas oaks dominated on shale (Foti

and Glenn 199 1). Shortleaf dominated southerly aspects and

intermediate slopes and, unlike today, was common on

northwest aspects. Black oak dominated west slopes and

white oak on north and east aspects. Post oak was more

important on gentle slopes and flats, whereas it is currently

found on steep slopes. Forests were generally more open

then than today, presumably due to more frequent fire (Foti

and Glenn 1991).

According to historical accounts and General Land O&e

surveys at the time of settlement, the vegetation mosaic of

the Ozark Mountains is relatively unchanged with the

exception of prairies, which have declined dramatically

(Tucker 199 1). While the amount of pine in the “original”

forest is still open to question, it is generally agreed that

pine was widespread in the region and increased southward.

Early surveyors described tornado damage but did not

mention fire and insect outbreaks. Tucker indicates that

understory vegetation is greater now than in the past, and

that mesic species [e.g., Acer saccharinum L.-A. jloridanum

(Chapm.) Pax complex] have expanded to drier areas in

more recent times, which may be related to reduced fire

frequency.

Based on fire scars from the period 1780-1938, mean fue

frequency across all sites in the Hot Springs NP was

estimated to be about 32 years and less on south and west

slopes (Johnson and Schnell 1985). Low-intensity fires often

fail to produce fire scars, which means that fire may have

occurred more often than these scars indicate. For the period

1788-1817, Foti and Glenn (1991) estimated a mean fire

interval of 7.25 years, also based on fire scars. Fire is

required for the maintenance of pine and oak-pine types in

the Ouachitas (Fountain and Sweeney 1985, Johnson and

Schnelll985, Fountain 199 1).

Appalachian Mountains: Blue Ridge

Distribution and composition-The Blue Ridge Mountains

extend from southern Pennsylvania to northern Georgia.

Braun (1950) describes the Southern Appalachian oakchestnut forest region as composed of oak-chestnut forests

on the slopes, mixed mesophytic forests in the coves, and

pine or oak-pine communities on dry slopes and ridges. The

demise of American chestnut earlier this century greatly

influenced forest structure and composition in this region,

Chestnut was dominant on intermediate to dry sites at a wide

range of elevations from 1,500 to 4,500 feet (457 to 1372

m) (Braun 1950). The most common replacement species

are chestnut oak, red maple, and northern red oak (Woods

and Shanks 1959, Golden 1974); and on drier sites, pitch

pine and scarlet oak. Historically, chestnut was a component

of the dry and dry-mesic oak-pine type.

The principal yellow pines of the Appalachians are variously

tolerant of fire. Cone serotiny in Table Mountain pine gives

it an advantage over shortleaf, pitch, and Virginia pines

when subjected to crown fues.6 Pitch pine has some degree

of cone serotiny (Fowells 1968), a characteristic shared, in

part, by shortleaf, and is more fire resistant than Virginia or

Table Mountain pine due to thicker bark and dormant buds

along the bole (Zobel 1969). Pitch and shortleaf pines

develop a basal crook in seedlings, which also may protect

them from fire (Little and Mergen 1966). Pitch pine is

6 Sanders, G., and E. Buckner. 1988. Fire: is it an essential component in

Table Mountain pine ecosystems? Poster session at 1988 National Society

of American Foresby Meeting, Rochester, NY.

8
typical of ecosystems that experience high-intensity fnes at

intervals of 10 to 30 years (Christensen 198 1).

White pine is not considered a principal pine species of the

dry and dry-mesic oak-pine type in this region, even though

it is a common associate of the oak-pine type. White pine

differs from the fire resilient yellow pine species mentioned

above in that it is somewhat more shade tolerant and is

commonly found on mesic sites (McCune 1988). White pine

distribution has expanded to upland sites in the Southern

Appalachians, likely as a result of forest disturbances in the

late 1800’s and early 1900’s (logging, farm abandonment,

severe fues), followed by a period of little or no fire. This

species may be a component of old-growth oak-pine forests

in the future since relatively old trees (>70 years) are

common on a variety of sites in the Southern

Appalachians.’ 8 It is likely that the fire regime before 1900,

including fires associated with Native Americans, had a

frequency that largely restricted white pine to more mesic

sites in the Appalachians, but this is not certain.

In the Great Smoky Mountains National Park (Great Smoky

Mountains NP), the oak-pine type is found in the ecotone

between xeric pine forests and intermediate oak forests and

often occurs on soils with a relatively high clay content, i.e.,

30 percent (Golden 1974, 1981; Callaway et al. 1987).

Topography and elevation were found to be the most

important factors affecting distribution of plant communities

(Whittaker 1956, Golden 1981, Callaway et al. 1987).

Whittaker (1956) described xeric pine forests and

intermediate oak forests of the Great Smoky Mountains NP

but did not describe the ecotonal “oak-pine” as a distinct

type. Descriptions for xeric pine forests and chestnut oakchestnut heath provide insight into the composition of oakpine forests (below and appendix A).

Southwest slopes and ridges in the Great Smoky Mountains

NP were occupied by Virginia pine below 2,500 feet (762

m), pitch pine heath between 2,200 and 3,200 feet (671 and

976 m), and Table Mountain pine heath between 3,500 and

4,500 feet (1067 and 1372 m). Intermediate oak forests

found adjacent to the xeric pine stands included red oakchestnut forests and the subxeric, chestnut oak-chestnut

heath. The latter occurs on open slopes under 3,700 feet

(1128 m), except south and southwest slopes. The red oak-

’ Carlson, P.J. 1995. An assessment of the old-growth forest resource on

National Forest System lands in the Chattooga River Watershed. Report to

Chattooga Ecosystem Demonstration Management Project, U.S.

Department of Agriculture, Forest Service, Region 8, Atlanta, GA. 83 p.

On tile with: USDA Forest Service, Southern Region, 1720 Peachtree

Road, NW, Atlanta, GA 30367.

* Continuous Inventory of Stand Condition (CISC) data files from the

following national forests: Pisgah, Nantahala, Chattahoochee, Sumter, and

Cherokee. On tile with: USDA Forest Service, Southern Region, 1720

Peachtree Road, NW, Atlanta, GA 30367.

chestnut type occupies sheltered south slopes and open

slopes above 3,700 feet (1128 m). Chestnut oak, American

chestnut, blackgum, sourwood, and red maple were

important associates in all three pine types, whereas scarlet

oak was an associate in the Virginia and pitch pine types.

Black, blackjack, and white oaks were important in the

Virginia pine community, whereas black locust and sassafras

were common in the Table Mountain pine heath.

Shrub importance was greater in the pitch and Table

Mountain pine types. Mountain laurel, trailing arbutus,

blueberry, deerberry, and huckleberry were common to all

three types. Of the lowbush blueberry species, Vaccinium

vacillans Torr. was common in Virginia and pitch pine

types, F. hirsutum Buckley in the pitch and Table Mountain

pine types (along with male blueberry) and K pallidurn

Aiton in the Table Mountain pine type. Highbush blueberry

(V. constablaei Gray), flame azalea, and fetterbush were

also common in the Table Mountain type. Herb species

common to all three types were little bluestem, galax, and

bracken fem. Wintergreen was common to the two higher

elevation forest types. Composition of xeric pine and oakheath forests is similar to oak-pine communities elsewhere

in the Southern Appalachians (appendix A).

In the Nantahala National Forest in North Carolina, McNab

and Browning ( 1993) identified a xeric, pine-oak heath on

steep, narrow, convex ridges with shallow soils, centered

around 3,500 feet (106 km). Scarlet and chestnut oaks with

sourwood and pitch pine dominated these relatively exposed

sites. Mountain laurel dominated the understory, whereas

low, deciduous heath (blueberry, huckleberry, and male

blueberry) was abundant on slightly more mesic sites. A dry

community of chestnut oak and red maple was adjacent to

the xeric community and occurred on steep sideslopes with

variable aspects and deeper soils. Low, deciduous heath or

tree seedlings comprised the understory.

Other studies clarifying the distribution of the oak-pine type

in the Appalachians include several that are descriptive of

river gorges found on the southeastern edge of the

Appalachians along the Blue Ridge Escarpment (Cooper

1963, Rodgers 1965, Mowbray 1966, Racine 1966,

Mowbray and Oosting 1968, Dumond 1970, Tobe et al.

1992). From many of these studies, Mowbray and Oosting

(1968) identified three intermediate to xeric types: (1)

chestnut oak-mixed oak on cool, moist slopes dominated by

chestnut, black, and northern red oaks; (2) oak-hickory on

drier west- and south-facing slopes dominated by scarlet and

chestnut oaks and several hickories; and (3) ridgetop

communities ranging from pure oak (scarlet and chestnut) to

pure pine (shortleaf, Virginia, pitch, and Table Mountain).

Associated with scarlet and chestnut oaks and pitch pine

were sourwood and red maple with a dense cover of

mountain laurel, lowbush bluebeny, deerberry, and bear

9
huckleberry. Mountain laurel cover decreased and blueberry

cover increased along the gradient from chestnut oak-heath

to pine leads and ridges.

Southern red, post, and blackjack oaks-typical Piedmont

species-and Virginia pine were described by Mowbray

(1966) as reaching their thermal limit at 1,800 to 1,900 feet

(549 to 579 m) in the Bearwallow Gorge, whereas Dumond

(1970) described the same three oak species as occurring

below 3,000 feet (9 15 m) in the Chattooga River Gorge.

Racine (1966) described the altitudinal limit of Virginia and

shortleaf pines as 2,800 feet (854 m), an elevation where the

pitch pine and scarlet oak association develops. Species

composition described for the Thompson River Gorge

(Racine 1966) is representative of most of the other

escarpment sites and is included in appendix A.

Shortleaf pine is an important species at lower elevations of

the Appalachians. In the Chauga River Gorge, shortleaf pine

dominates the ridges below 2,000 feet (610 m), whereas a

rare Table Mountain pine-Virginia pine association is found

above 2,000 feet (610 m) (Tobe et al. 1992). Shortleaf type

associates include red maple, dogwood, blackgum,

sourwood, Table Mountain pine, Virginia pine, pitch pine,

scarlet and southern red oaks, with lowbush bluebeny and

storax in the shrub layer (appendix A). The chestnut oak

type, the most common in this gorge, merges with pine types

on north and east aspects of the upper slopes. Scarlet oak is

a consistent codominant in the chestnut oak and pinedominated forests of the upper gorge slopes.

In a central Appalachian gorge in Virginia, Ramsey et al.

(1993) described an oak-pine heath on thin, acid soils on

south and west slopes, dominated by Virginia, pitch, and

Table Mountain pines, and northern red, chestnut, and

scarlet oaks. The understory included bear oak, serviceberry

[Amelunchier arborea (Michx. f.) Fem.], mountain laurel,

chestnut sprouts, sweet-fern [Comptonia peregrinu (L.)

Coult.], lowbush blueberry, deerberry, huckleberry, minniebush [Menziesiu pilosu (Michx.) Juss.], wild azalea,

Catawba rhododendron, spirea (Spirea betulifoliu Pallas),

buffalo nut, hawthome, and sassafras.

Old-growth forest examples-Few old-growth oak-pine

stands have been studied or documented in this region. At

2,200 to 3,000 feet (67 1 to 9 15 m) in the Scarlet Oak

Natural Area of the Pisgah National Forest in North

Carolina, oak-pine stands considered high in “virgin

attributes” were described by Delapp and Wentworth’

(appendix A). Scarlet and chestnut oaks, sourwood, pitch

pine, and red maple dominated on broad, south-facing leads

and knobs. Other species included mountain laurel,

9 Delapp, J.A., and T.R. Wentworth. 1977. Proposed research natural areas

in the Southern Appalachian Mountains. 117 p. Unpublished report. On

tile with: Highlands Biological Station, Highlands, NC 28741.

huckleberry, lowbush blueberry, deerberry, Smilux species,

pipsissewa, and bracken fem. Pitch pine increased in

importance on drier sites in these scarlet oak-dominated

stands. Scarlet oak was considered the “climax” species on

these sites.

The remaining descriptions of old growth are more general

and do not provide detailed, stand-level information.

Carlson (see footnote 7) assessed old growth across parts of

three national forests. The oak-pine type was

underrepresented as old growth as compared to its current

relative abundance in younger age classes. Carlson

described stands containing pitch and shortleaf pines; white,

black, and scarlet oaks; and large red maple and white pine.

The yellow pine species exceeded 200 years in age.

Although some of these stands approximated old-growth

conditions, the effects of chestnut blight, early logging and

fire, and 20”-century fire suppression greatly influenced

current stand composition and structure.

Zahner” described dry oak and oak-pine old-growth types

for the Highlands Ranger District on the Nantahala National

Forest in North Carolina as occurring on high ridges and

upper, south-facing slopes. Species included white oak,

scarlet oak, black oak, chestnut oak, pitch pine, white pine,

witch-hazel (Humumelis virginiunu L.), black locust, Table

Mountain pine with Catawba rhododendron, mountain

laurel, and mountain myrtle [Leiphyllum buxifolium (Berg.)

Ell.] in the understory.

Yost et al. (1994) used five forest types to delineate old

growth in the Great Smoky Mountains NP. Their subxeric

and xeric oak types contained yellow pine species and

corresponded closely with the dry and dry-mesic oak-pine

type. Scarlet, chestnut, and black oaks dominated the

subxeric type, with an understory dominated by mountain

laurel, huckleberry, and Rhododendron species. Xeric sites

were dominated by blackjack, scarlet, and chestnut oaks,

with Virginia, pitch, and Table Mountain pine, sourwood,

blackgum, and red maple as associates and mountain laurel

and blueberry species in the understory. Their assessment of

these dry types for the Great Smoky Mountains NP is

incomplete at this time.

Presettlement forests and disturbance-The abundance of

the oak-pine type in the Appalachians at the time of

European settlement is not clear. Bartram (1958) described

the vegetation in the Carolina mountains but makes only one

reference to pines. He mentioned white pine, Scotch pine

I0 Zahner, R. 1990. Upland hardwood and pine-hardwood forest of the

Southern Appalachian Mountains-old-growth stands: definitions and

characteristics for identification, inventory, and restoration on the

Highlands Ranger District, Nantahala National Forest. 13 p. Unpublished

report. On tile with USDA Forest Service, Highlands Ranger District,

Highlands, NC 2874 1.

10
(Pinus sylvestris L.), and other species at a South Carolina

site. His reference to Scotch pine may be an incorrect

identification of Virginia or Table Mountain pine since

Scotch pine is not native to the area. Sargent (1884)

describes the North Carolina and Virginia mountains as

containing “northern pines” but focused generally on the

“great body of relatively undisturbed hardwood forest” of

that region. Malter (1977) quotes a 190 1 article describing

the forests of the Citico Creek in Tennessee before the

extensive logging of the early 1900’s: “the finest timber

lands he has ever seen, barring none. Pine, oak, poplar and

hemlock abound. . . and the forest affords uniformly good

timber throughout its entire length and breadth, speaking

with respect to the 43,000 acres of land owned or controlled

by the company.”

The pollen record indicates the group of southern yellow

pine species expanded northward into the Southern

Appalachians approximately 10,000 years ago as glaciers

retreated (Delcourt and Delcourt 1987) at approximately the

same time Native Americans entered the region. Wind and

ice storm damage and insect outbreaks, coupled with fire,

could have resulted in large burns in the Appalachians. Use

of fire by Native Americans (Buckner 1989, Van Lear and

Waldrop 1989), coupled with lightning fire and other natural

disturbances, probably enhanced and maintained the oakpine type in presettlement times.

Although it is widely acknowledged that Native American

land use strongly impacted southeastern landscapes, the

magnitude and frequency of their disturbances across large

regions is uncertain. It is our view that natural disturbances

(wind and ice storms, insects, pathogens, and lightning fire)

may have been sufficient to maintain the oak-pine type in

the Appalachians. The assessment by Barden and Woods

(1974, 1976) that lightning-caused crown fires are extremely

uncommon in the Appalachians should be interpreted

cautiously. The short duration of their study and the fact that

the lightning fires they investigated were successfnlly

extinguished (except those that went out on their own) rather

than having been allowed to bum unhindered or to smolder

until burning conditions improved, allow other

interpretations. Frost (1995) suggests the importance of

lightning fire on exposed ridges and upper slopes in the

Appalachians in maintaining pine and pine-oak types. Frost

estimated presettlement fire frequency (for the most exposed

parts of landscape) to be 7 to 12 years in the lower

mountains [13,000 feet (915 m)] and more than 12 years in

higher mountains. Frost suggests the Indian fires were of

greater impact in more topographically diverse areas such as

the mountains, upper Piedmont, and sandhills compared to

the less dissected Piedmont and Coastal Plain. Whittaker

(1956) viewed pine stands and oak-chestnut heaths as selfmaintaining, whereas more recent studies have shown that

this is, not the case, except possibly on extremely dry sites

(Barden 1976, Williams and Johnson 1992).

Old-growth representatives of the dry and dry-mesic oakpine type are uncommon in the Appalachians. However,

younger stands are common and often exist in the ecotone

between xeric pine and intermediate to dry oak-dominated

forests. The prevalence of this type is due to fire and other

disturbances. Scarlet oak and chestnut oak are the most

common oak species of this type, whereas the relative

dominance of the four yellow pine species is determined by

elevation, exposure, and the type and magnitude of

disturbance.

Appalachian Mountains: Plateau

Distribution and composition-The Appalachian Plateau

extends from central New York to southwestern Alabama.

We describe the vegetation in the unglaciated sections from

central Pennsylvania southward. Shimer (1972) described

this region’s appearance from the air as a choppy sea of

small mountains oriented randomly, regionally appearing as

a basin-type structure with cuesta scarps along its border.

On its eastern front are the Allegheny and Cumberland

Mountains, with the majority of the region consisting of the

Allegheny and Cumberland Plateaus. Braun (1950)

described four sections within these plateaus: (1) rugged

eastern area-occurring mostly between the Allegheny and

Cumberland Mountains; (2) low hills belt-comprising most

of West Virginia, northeastern Kentucky, and southern

Ohio; (3) knobs border area-consisting of a series of

conical hills at the western edge of the Allegheny Plateau in

northern Kentucky and southern Ohio; and (4) cliff

section-extending from Berea, KY, to Alabama, initially

along the western plateau border as a distinct escarpment but

widening in Tennessee and Alabama where it forms most of

the plateau.

Although Braun (1950) described the overall area as the

mixed mesophytic forest region, other types such as oak,

oak-pine, and pine types were represented on exposed areas

throughout the region. In the Cumberland Mountains (Braun

1935, 1942, 1950), dry pine and oak forests occupied dry

ridges and upper south slopes. Virginia, pitch, and shortleaf

pines occurred with chestnut; black, white, and scarlet oaks;

blackgum; American chestnut; red maple; yellow-poplar;

dogwood; sassafras; and sourwood. The oak-pine forests

were small in stature on the shallow soils, increasing in size

as soil became deeper, grading into chestnut oak and

American chestnut communities.

Braun (1935) described several oak-pine to pine heath

stands that she considered representative of “primary”

forests on Pine Mountain in Kentucky. Overstory

composition was described as above. Understory shrubs

included mountain laurel, lowbush blueberry, deerbexry, and

others. See appendix A for additional species.

11
Yellow pines were apparently not as prevalent in the

Allegheny Mountains as they were in the Cumberland

Mountains based on Braun’s (1950) descriptions but were

components of the oak-chestnut type on upper slopes and

ridges. Braun cites Jennings’ (1927) description of a

chestnut-chestnut oak-black oak association occurring with

pitch pine on the upper slopes and “undisturbed top” of

Chestnut Ridge in Pennsylvania. Understory species

included mountain laurel, azalea, blueberry, trailing arbutus,

and wintergreen.

Within the rugged eastern area of the Cumberland and

Allegheny Plateau region, several observers have described

the oak-pine type. Braun (1950) describes pitch and

shortleaf pines as scattered along ridges and occasionally in

small groups in oak-chestnut and oak-hickory communities.

Martin and Shepherd (1973) described upland tree and

shrub species in the “virgin” portion of the Lilley Comett

Woods in Kentucky that included pitch, Virginia, and

shortleaf occurring occasionally. In the nearby Robinson

Forest, shortleaf and pitch pines were minor components of

upper slope, second-growth forests (Phillippi and Boebinger

1986).

In the low hills section, Braun cites Crandall’s (1876)

description of the “original” vegetation of an area in eastern

Kentucky where pine species accounted for 11 percent of

the composition on hilltops dominated by chestnut oak.

Overall, pine was likely a minor component of the oak- and

oak-hickory-dominated slopes and ridges of the low hills

section.

The conical hills of the narrow knobs border area are

underlain by shales, limestones, and sandstones. Oak,

chestnut, oak-hickory, and pine types predominated near the

summit of ridges or knobs (Braun 1950). Pines capped the

knobs and extended down the steep, southwestern slopes.

The heath layer is similar to that described for the

Cumberland and Allegheny Mountains. The pine-oak and

chestnut heath type communities dominated on

noncalcareous soils (sandstone or shale), whereas oakhickory was more common on limestone ridgetops.

The cliff section of the Cumberland Plateau in southern

Kentucky is flat to rolling with shallower drainages and

gorges than the area farther south. The plateau supports oakhickory and oak-pine forests of white, black, and post oaks;

mockemut and shagbark hickories [C. ovutu (P. Mill.) K.

Koch]; yellow-poplar; dogwood; shortleaf and pitch pines;

and often blackgum, red maple, and Virginia pine (Braun

1950). Lowbush blueberry, wild azalea, sawbrier, grape, and

box huckleberry [ Guylus.suciu brachyceru (Michx.) Gray]

make up the understory. Toward the western margin of the

plateau along the western escarpment, sandstone cliffs

become prominent. Forests there vary from open pine woods

with grassy understories to pine and pine-oak heath with

scarlet and chestnut oaks and an understory dominated by

mountain laurel, lowbush blueberry, huckleberry, or

occasionally, box huckleberry (Braun 1950).

The southern part of the cliff section in Alabama and

Tennessee consists of oak, oak-hickory, and oak-pine

communities (Braun 1950). At the plateau’s margin above

the gorges, pine heath communities dominate. Hinkle (1989)

noted Virginia pine occupying the dry ridges and cliff edges

and, on deeper soils, identified a shortleafpine-white oak

type that he viewed as succeeding white oak. He refers to

this section in Tennessee as the plateau uplands.

Schmalzer (1988) described three moderately xeric oak-pine

or pine communities of the Obed River Gorge in Tennessee

as white pine-white oak-chestnut oak, white oak-scarlet oak,

and Virginia pine. Virginia pine was common and persistent

on shallow siliceous soils of cliff edges and on upland sites

following logging and fire. Shortleaf, Virginia, and white

pines were significant components of the white oak-scarlet

oak type. Unlike other areas of the Appalachian Plateau,

chestnut and hickory were not significant components of

oak-dominated upland plateau communities in Tennessee.

The prevalence of fire may explain the scarcity of hickory.

Schmalzer (1988) and Hinkle (1989) acknowledge the role

of fire in affecting the composition of these upland types.

Old-growth forest examples-No documented examples

of old-growth oak-pine exist to our knowledge.

Presettlement forests and disturbance--Little information

is available on presettlement distribution of the oak-pine

type in the region other than that described above. In the

plateau region of Alabama, Harper (1943) described

shortleaf pine and mixed oaks as occurring in dry woods

around Little Mountain in the Tennessee Valley and coal

region. Both loblolly and shortleaf were significant

components of the “original” forest of the latter, occurring

with post, southern red, and white oaks. In the Tennessee

River Valley as a whole, pine was a minor component

(Mohr 1901, Harper 1943).

On the Alabama tablelands of the Warrior and Coosa

Basins, Mohr (190 1) described chestnut oak as a dominant

species, occurring with post and southern red oaks, or,

rarely, blackjack, black, white, and scarlet oaks. Shortleaf

was more prevalent on lower elevation ridges with thinner

soils and was rare when chestnut oak was dominant. He

found dense groves of loblolly pine growing in poorly

drained depressions and Virginia pine, on the poorest sites,

mostly above 1,200 feet (366 m). Lowbush blueberry,

deerbeny, and wild azalea were important understory

species associated with the drier oak and pine types.

In summary, mixed oak and pine communities containing

pitch, shortleaf and Virginia pines, or both, are prevalent in

1 2
the Cumberland Mountains and southern portion of the

Appalachian Plateau, often growing on sandstone or shalederived soils on exposed sites. Composition of these

communities does not vary significantly across the region.

Historically, fire has been an important factor in maintaining

the pine and oak-pine types on these droughty uplands.

Appalachian Mountains: Valley and Ridge

Distribution and composition-The Valley and Ridge

province extends from the St. Lawrence River lowland to

northeast Alabama (Shimer 1972). The section in northern

Pennsylvania and from New Jersey northward was glaciated

and is mostly beyond the range of the dry and dry-mesic

oak-pine type. This province consists of long ridges, mostly

oriented southwest to northeast, predominately sandstone

and conglomerate, with the intervening valleys or lowlands

underlain by shale and limestone. The Great Valley or

Appalachian Valley, a series of large valleys, occurs

between the easternmost ridge of the province and the Blue

Ridge. Braun (1950) noted that most of these ridges were

not high enough to have well-defined ahitudinal variations

in vegetation. Because ridge forests are easily accessible,

most have been logged heavily by the charcoal, railroad, and

chemical industries.

In the glaciated areas of the Valley and Ridge region, dry to

xeric pitch pine-oak communities with heath understories

have been found in northern New Jersey (Niering 1953) and

southeastern New York (McIntosh 1959). These

communities were similar to those of the New Jersey Pine

Barrens (Buell and Cantlon 1950), except for different

species of Gaylussacia and Vaccinium. Many of these

forests are stunted and considered too xeric for the dry and

dry-mesic oak-pine type, although Niering does describe 17-

to 19-inch (43- to 48-cm) d.b.h. pitch pines 140 years old in

a pine-oak stand.

Braun (1950) considered the tmglaciated parts of the Valley

and Ridge to be in the oak-chestnut region except for the

southern end that includes the Coosa Valley in Alabama,

which she considered part of the oak-pine region. She

described forests of the ridges and mountain ranges for three

areas. The northern area, in Pennsylvania, included a scarlet

oak or scarlet oak-pine community on dry, south, and west

slopes. The second area, from Spruce Knob, WV, south 100

miles (160 km), had chestnut oak dominating dry, upper

slopes with pines common on the cliff margins. Chestnut

oak, northern red oak, and Virginia pine occupied open

shale barrens. The third area, in southern Virginia and

Tennessee, was primarily hardwood. The region from

Knoxville, TN, south was discussed by Braun in the context

of valley floor vegetation and lacked mountain ranges.

White oak was dominant in the valley, whereas on low shale

ridges, oak communities (white, black, scarlet, and chestnut)

were prevalent. White, pitch, and Virginia pines were

associates in the north, although southern pines occurred in

the south with blackjack and southern red oaks. Martin

(197 1) also describes the prevalence of white oak

communities in eastern Tennessee, where white oak is often

codominant with other oaks and Virginia and shortleaf

pines.

In the oak-pine forests of southwest Virginia, scarlet,

chestnut, and black oaks were dominant on side slopes,

whereas pitch and pitch pine-scarlet oak were dominant on

convex sites (McEvoy et al. 1980). Lowbush blueberry,

huckleberry, and mountain laurel were common on these

sites. Tree cover declined as shrub cover increased from

mesic to xeric sites, as was found in the Great Smoky

Mountains NP (Whittaker 1956). Elsewhere in southwest

Virginia, relatively undisturbed Table Mountain pine stands

occupied southwest slopes at 2,500 feet (762 m) and

contained scarlet and chestnut oaks, sassafras, and blackgum

(Williams and Johnson 1992). Within the Valley and Ridge,

Table Mountain pine typically occupies steep, lower, and

middle slopes, often on shale outcroppings (Zobel 1969) or

ridge-capping sandstones at higher elevations.

Old-growth forest examples-No documented examples

of old-growth oak-pine exist to our knowledge.

Presettlement forests and disturbance-Little is known

about the “original” forests of the Valley and Ridge

province (Clarkson 1966). Clarkson grouped the “original”

forest of the Monongahela National Forest, WV, into four

types. Mixed oak-chestnut was the only type that contained

yellow pines (primarily pitch, but also Table Mountain,

shortleaf, and Virginia). Pines were abundant on southern

and eastern slopes on thin soils derived from shale or

sandstone. Early forests of West Virginia were described by

Sargent (1884), who quotes a report from a Mr. Pringle,

who described the long mountain chains of West Virginia

after leaving Pemsylvania: “ . . . fewer evergreens appeared

than I had previously seen. A few slopes were principally

occupied by pine variety, but the mountains of this region

were covered with a growth of deciduous trees, white, black,

red, Spanish, and chestnut oaks, hickories, butternuts, black

walnuts, yellow-poplars, locusts, elms, sugar maples . . . .”

The presettlement forests in Pennsylvania were comprised

of oak (42 percent), pine (27 percent), and American

chestnut (13 percent) on ridges and oak (56 percent), pine

( 13 percent), and hickory ( 12 percent) in the valleys

(Nowacki and Abrams 1992). Fire and other disturbances

are thought to be required to maintain oak and yellow pine

in these forests. The pine component of these forests has

been drastically reduced by past logging activities.

Repetitive harvesting for charcoal production greatly

fostered vegetative species (i.e., oak, chestnut) at the

expense of pine. Due to recent tire suppression, composition

is shifting toward greater maples, black cherry, and black

1 3
birch (Betula lenta L.). Dry sites are currently dominated by

blueberry and huckleberry species with mountain laurel

confined to ridge sites. Northern red oak is an important

associate of chestnut oak on drier ridge sites, whereas scarlet

oak is an important associate of white oak on coarse

textured soils in the valley. The authors suggest that the

increased presence of scarlet oak in valleys more so than on

ridges may be a local phenomenon. They propose that the

abundance of northern red oak on ridges may have been due

to more presettlement fires in the valleys where most Native

Americans resided because northern red oak is less tolerant

of fire than most oaks (Lorimer 1985).

In the southern part of the Valley and Ridge province,

surveys of northwest Georgia from the early 1800’s indicate

that the oaks-to-pines-to-hickories ratio was 50: 18:8

(Plummer 1975). Post oak was the most abundant oak with

hickories and American chestnut on ridges. Elevation of this

section ranged from 600 to 1,700 feet (183 to 5 18 m), In the

Coosa Valley of Alabama, many of the chert hills and

sandstone ridges supported longleaf pine on sandy and

gravelly soil (Harper 1943). On dry soils, shortleaf pine and

southern red and post oaks were dominant, with dogwood,

rusty black-haw, and hawthome in the understory. Loblolly

was widely distributed. About 40 percent of the “original”

forest was estimated to have been evergreen (mostly pine).

An assessment of age structure and factors affecting

recruitment in relatively undisturbed Table Mountain pineoak communities of the Jefferson National Forest indicated

that Table Mountain pine was not self-maintaining and that

successful seedling recruitment and establishment were

unlikely without fire (Williams and Johnson 1992). In

general, without fire, pines become less abundant in these

forests and, in some cases, oaks eventually decrease as well

(Nowacki and Abrams 1992). The intensity of

postsettlement disturbance in this area conceals the

“original” abundance of pine or oak-pine types in the region,

since much of this disturbance has decreased pine

abundance.

Forest Composition: Synthesis across Regions

Species composition of the oak-pine forest type described

above is shown for several old-growth and other

representative stands in appendix A. Pinus, Quercus, Carya,

and Vaccinium were represented in all seven old-growth

stands. Herb and vine species found in most stands included

Carex, Panicum, broom-sedge, and other grasses,

pipsissewa, Desmodium, bracken fern, Smilax, Virginia

creeper, and Vitis. Shortleafpine and white oak were the

most widely distributed trees across all stands (appendix A).

Shortleaf was the primary pine species of the dry and drymesic oak-pine type in the Piedmont, Coastal Plain, and

Interior Highlands, whereas pitch was predominant in the

mountainous regions. Other widely distributed oaks include

black, post, southern red, and blackjack. Scarlet and

chestnut oaks were more prevalent in the Appalachian

stands. Red maple, blackgum, and sourwood were common

across most sites shown in appendix A and may be

increasing on most sites due to fire suppression. Dogwood

was common except in some of the mountain stands.

Vaccinium species (I’. vacillans and V. stamineum

primarily) were the most common shrubs across regions,

whereas mountain laurel was common on mountain sites.

Gaylussacia species j-primarily G. baccata (Wang.) K.

Koch] were common on half the sites.

Narrative of Old-Growth Conditions

In this section, we describe characteristics of the living and

dead components of old-growth stands found in the

Piedmont, Blue Ridge, Ouachita Mountains, and Gulf

Coastal Plain (table 2). See appendix B for additional

information on each of the old-growth stands presented in

table 2. The most complete old-growth attribute information

available for the dry and dry-mesic oak-pine type is from the

John de la Howe (JDLH) forest tract in the South Carolina

Piedmont. Since this stand will be frequently referred to in

this section, a short description of it follows.

The JDLH stand is an approximately 120-acre [48.6-hectare

(ha)], 200-year-old stand that in recent years has suffered

significant pine mortality. Prior to settlement, the general

area was subjected to aboriginal fires and likely consisted of

open woods with scattered prairie-like openings (Logan

1859, 1980). It is located in an area that was settled by

French Huguenots around 1770, abandoned in the 1790’s,

and has been generally protected since. From current soil

and vegetation conditions (age, composition) and from

archeological evidence (Steen 1993), site disturbance likely

included some land clearing for crops and timber as well as

woodland livestock grazing. The disturbances that preceded

abandonment in the late 1700’s resemble, in terms of their

effect on the landscape, relatively large-scale natural

disturbance (disease, wind, ice, tomados) followed by fne.

Scattered cutting in the periphery of the stand and pine

salvage in a few areas has taken place in recent years. These

areas were avoided when sampled. Data was collected at

four levels of mortality, which will be described later.

1 4
Table 2 (English units)--Standardized table of old-growth attributes for dry and dry-mesic oak-pine

forests

Quantifiable attribute Range’ Mean

Number

of standsb References

Stand density (no./acre)

John de la Howe tract

-all species 24 in. d.b.h.

Shortleaf and loblolly pines

Oak species

130-183 171

l-42 25

31-51 42

7-22 15

1-15 9

l-5 4

1 This study

-all species 220 in. d.b.h.

Shortleaf and loblolly pines

Oak species

Duke Forest 1 Oosting 1942

-all species tl in. d.b.h.c 134

Shortleaf pine 32

Oak species 93

Scarlet Oak Natural Area

-all species 23 in. d.b.h.

Pitch pine

Oak species

3 12-328 320

20-134 77

117-200 160

2 Delapp and Wentworth d

Lake Winona Research Natural Area

-all species 24 in. d.b.h.

Shortleaf pine

Hardwoods

1 Fountain 199 1

-shortleaf pine X!O in. d.b.h.

209

138

71

3

Roaring Branch Research Natural Area

-all species 24 in. d.b.h.

Shortleaf pine

Oak species

-shortleaf pine 220 in. d.b.h.

223-225

3&56

115-156

&3

224

42

137

2

2 Fountain and Sweeney 1985

Hot Springs National Park

-all species 3.5 in. d.b.h.

Shortleaf pine

Oak species

313-618 496

67-367 212

117-367 202

12 Dale and Watts 1980

Kisatchie National Forest

-all species 25 in. d.b.h.

Shortleaf and loblolly pines

Oak species

2 Martin and Smith 199 I

-all species 220 in. d.b.h.

Shortleaf and loblolly pines

Oak species

61-107 84

14-24 18

38-75 57

lo-16 13

610 8

4-6 5

Stand basal area (ft2/acre)

John de la Howe tract

-all species 24 in. d.b.h.

Shortleaf and loblolly pines

Oak species

1 This study

-all species 220 in. d.b.h.

Shortleaf and loblolly pines

Oak species

64150 118

5-80 47

20-43 35

22-7 1 47

5-55 28

4-18 13

Duke Forest

-all species 21 in. d.b.h.c

Shortleaf pine

Oak species

1 Qosting 1942

128

38

76

15
Table 2 (English units)-Standardized table of old-growth attributes for dry and dry-mesic oak-pine

forests (continued)

Quantifiable attribute Rangea Mean

Number

of standsb References

Stand basal area (&acre) (cont.)

Scarlet Oak Natural Area

-all species L3 in. d.b.h.

Pitch pine

Oak species

lake Winona Research Natural Area

-all species 24 in. d.b.h.

Shortleaf pine

Hardwoods

-shortleaf pine 220 in. d.b.h.

Roaring Branch Research Natural Area

-all species 24 in. d.b.h.

Shortleaf pine

Oak species

-shortleaf pine 20 in. d.b.h.

Hot Springs National Park

-all species 23.5 in. d.b.h.

Shortleaf pine

Qak species

Kisatchie National Forest

-all species t5 in. d.b.h.

Shortleaf and loblolly pines

Oak species

-all species X0 in. d.b.h.

Shortleaf and loblolly pines

Oak species

Age of large trees (yrs) ’

John de la Howe tract

Shortleaf pine

Loblolly pine

White oak

Hickory speciesf

Post oak

Southern red oak

Yellow-poplar

Scarlet Oak Natural Area

Pitch pine

Scarlet oak

Chestnut oak

Great Smoky Mountains National Park

(max. age)

Chestnut oak

White oak

Scarlet oak

Black oak

Pignut hickory

Pitch pine

Table Mountain pine

Virginia pine

90-l 10 100

l&57 33

36-68 52

80-81 81

2430 27

4246 44

&9 5

57-l 12

1860

2&55

79

34

34

59-128 94

27-45 36

30-81 56

33-50 42

22-33 28

11-17 14

89-205 144

79-189 117

82-207 149

142-207 179

101-216 144

64-190 124

94-194 160

117-163

73-123

66170

347

344

165

180

327

148

204

135

2 Delapp and Wentworth d

1 Fountain 199 1

12 Dale and Watts 1980

Fountain and Sweeney 1985

Martin and Smith 199 1

This study

This study

Blozan 1994

16
Table 2 (English units)--Standardized table of old-growth attributes for dry and dry-mesic oak-pine

forests (continued)

Quantifiable attribute Range* Mean

Number

of standsb References

Age of large trees (yrs) e (cont.)

Chattooga River Watershed

(max. age)

Chestnut oak

White oak

Scarlet oak

Black oak

Mockernut hickory

Pitch pine

Shot-deaf pine

Virginia pine

Lake Winona Research Natural Area

Shortleaf pine

Roaring Branch Research Natural Area

Shortleaf pine

Hot Springs National Park

Shortleaf pine

Number of 4-in size classes ’

John de la Howe tract

Shortleaf pine

Hardwoods

362

348

190

211

335

235

300

108

108-314

133-275

124-247

8

7

Carlson 1 995g

Stahle et al. 1985

Stahle et al. 1985

Stahle et al. 1985

This study

Scarlet Oak Natural Area

Pitch pine

Hardwoods

2

Lake Winona Research Natural Area

Roaring Branch Research Natural Area

Shortleaf pine

Hardwoods

Kisatchie National Forest

Shortleaf and loblolly pines

Hardwoods

D.b.h. of large trees (in.) e

John de la Howe tract

Shortleaf pine

Loblolly pine

White oak

Hickory specie?

Post oak

Southern red oak

Yellow-poplar

6

4

I

5

1 l-32

12-39

12-33

lo-23

8-19

12-32

12-31

19

21

20

16

14

22

22

Fountain 199 1

Fountain and Sweeney 1985

Martin and Smith 1991

This study

Roaring Branch Research Natural Area

Shortleaf pine

Standing coarse woody debris (no.lacre)

John de la Howe tract

Snags 24 in. d.b.h.

Snags a0 in. d.b.h.

426 19

5-69 33

2-16 8

1 Stahle et al. 1985

1 This study

17
Table 2 (English units)-Standardized table of old-growth attributes for dry and dry-mesic oak-pine

forests (continued)

Quantifiable attribute Rangea Mean

Number

of standsb References

Standing coarse woody debris (no./acre)

(cont.)

Hot Springs National Park i

(mass only-tons/acre)

Snags >3 in. d.b.h.

Downed coarse woody debris

John de la Howe tract

Volume (ft3/acre)

Mass (tons/acre)

3 Johnson and Schnell 1985

1.9-13.1 8.7

1 This study

147-2528 1.545

7.4-25.4 15.4

Hot Springs National Park i

Mass (tons/acre)

3 Johnson and Schnell 1985

.94.2 3.1

Tree canopy structure

John de la Howe tract

Number of canopy layers (l-3)

Percent canopy in gaps

Mean gap size’ (acres)

2-3

24-80

.002-SO3

31

.06

1 This study

a For the John de la Howe tract, this represents the range of means from areas showing different degrees of mortality, and applies to all

attribute data except age, diameter, and gap size.

b Number of stands included (may not equal number of citations).

’ Oosting did not define the diameter limit for overstory. Density values are for overstoty only. If understory is included, density of trees

tl in. in d.b.h. is 687,89, and 154 for all trees, pines, and oaks, respectively. Basal area values are given for all trees 21 in. in d.b.h.

d Delapp, J.A., and T.R. Wentworth. 1977. Proposed research natural areas in the Southern Appalachian Mountains. 117 p. Unpublished

report. On file with: Highlands Biological Station, Highlands, NC 2874 1.

e Includes dominant and codominant trees that comprise the upper canopy. Mean diameter for trees from the John de la Howe tract is the

quadratic mean.

f

Includes mockemut, shagbark [&vu ovatu (Mill.) K. Koch], and pignut hickory species.

g Carlson, P.J. 1995. An assessment of the old-growth forest resource on National Forest System lands in the Chattooga River Watershed.

Report to Chattooga Ecosystem Demonstration Management Project, U.S. Department of Agriculture, Forest Service, Region 8, Atlanta,

GA. 83 p. On file with: USDA Forest Service, Southern Region, 1720 Peachtree Road, NW, Atlanta, GA 30367.

’ For inclusion, 1 percent of all trees must be present within a size class-size classes start at 4 in.

‘Average across three stands: (1) stand dominated by medium to large, actively growing shortleaf, (2) a “decadent” old-growth shortleafdominated stand, and (3) an old-growth oak-hickory stand with a few large shortleaf pines.

’ Refers to canopy gaps <l acre in size. Gaps exceeding 1 acre were variable sized patches of extensive pine mortality that accounted for

58 percent of the total gap area.

18
Table 2 (metric units)-Standardized table of old-growth attributes for dry and dry-mesic oak-pine

forests

Quantifiable attribute Rangea MeG%l

Number

of stand.? References

Stand density (no./acre)

John de la Howe tract

-all species 210 cm d.b.h.

Shortleaf and loblolly pines

Oak species

-all species 250.8 cm d.b.h.

Shortleaf and loblolly pines

Oak species

321-452 422

2-104 62

77-126 104

17-54 37

2-37 22

2-12 10

Duke Forest

-all species 22.5 cm d.b.h.c 331

Shottleaf pine 79

Oak species 230

Scarlet Oak Natural Area

-all species 27.6 cm d.b.h.

Pitch pine

Oak species

771-810 790

50-330 190

289-494 395

Lake Winona Research Natural Area

-all species 210 cm d.b.h.

Shortleaf pine

Hardwoods

-shortleaf pine 250.8 cm d.b.h.

516

341

175

7

Roaring Branch Research Natural Area

-all species 210 cm d.b.h.

Shortleaf pine

Oak species

-shortleaf pine 250.8 cm d.b.h.

551-556

74-138

284-385

o-7

2

553

104

338

5

Hot Springs National Park

-all species 28.9 cm d.b.h.

Shortleaf pine

Oak species

773-1526

165-906

289-906

12

1225

524

499

Kisatchie National Forest

--all species 212.7 cm d.b.h.

Shortleaf and loblolly pines

Oak species

-all species 250.8 cm d.b.h.

Shortleaf and loblolly pines

Oak species

151-264

35-59

94-185

25-40

15-25

l&15

2

207

44

141

32

20

12

Stand basal area (m2/ha)

John de la Howe tract

-all species 210 cm d.b.h.

Shortleaf and loblolly pinesOak species

-all species 250.8 cm d.b.h.

Shortleaf and loblolly pines

Oak species

15-34 27

1-18 1 1

5-10 8

5-16 1 1

l-13 6

l-4 3

Duke Forest

-all species 22.5 cm d.b.h.c

Shortleaf pine

Oak species

29

9

17

Oosting 1942

1 This study

Delapp and Wentworth d

Fountain 1991

Fountain and Sweeney 1985

Dale and Watts 1980

Martin and Smith 199 1

This study

1 Oosting 1942

19
Table 2 (metric units)-Standardized table of old-growth attributes for dry and dry-mesic oak-pine

forests (continued)

Quantifiable attribute Range’ Mean

Number

of standsb References

Stand basal area (m*/ha) (cont.)

Scarlet Oak Natural Area

-all species 27.6 cm d.b.h.

Pitch pine

Oak species

Lake Winona Research Natural Area

-all species 210 cm d.b.h.

Shortleaf pine

Hardwoods

-shortleaf pine 250.8 cm d.b.h.

Roaring Branch Research Natural Area

-all species 210 cm d.b.h.

Shortleaf pine

Oak species

-shortleaf pine 250.8 cm d.b.h.

Hot Springs National Park

-all species 27.6 cm d.b.h.

Shortleaf pine

Oak species

Kisatchie National Forest

-all species 212.7 cm d.b.h.

Shortleaf and loblolly pines

Oak species

-all species 250.8 cm d.b.h.

Shortleaf and loblolly pines

Oak species

Age of large trees (yrs) e

John de la Howe tract

Shortleaf pine

LobJolly pine

White oak

Hickory species’

Post oak

Southern red oak

Yellow-poplar

Scarlet Oak Natural Area

Pitch pine

Scarlet oak

Chestnut oak

Great Smoky Mountains National Park

(max. age)

Chestnut oak

White oak

Scarlet oak

Black oak

Pignut hickory

Pitch pine

Table Mountain pine

Virginia pine

21-25 23

2-13 8

8-16 12

18-19 19

6-7 6

10-I I 10

o-2 1

13-26 18

414 8

5-13 8

14-29 22

610 8

7-19 13

8-11 10

5-8 6

3 4 3

89-205 144

79-l 89 117

82-207 149

142-207 179

101-216 144

64-190 124

94-194 160

2 Delapp and Wentworth ’

Fountain 199 1

26

23

3

0

2 Fountain and Sweeney 1985

12 Dale and Watts 1980

2 Martin and Smith 199 1

1 This study

1 This study

117-163

733123

66170

Blozan 1994

347

344

165

180

327

148

204

135

20
Table 2 (metric units)-Standardized table of old-growth attributes for dry and dry-mesic oak-pine

forests (continued)

Quantifiable attribute Range’ Mean

Number

of standsb References

Age of large trees (yrs) e

(cont.)

Chattooga River Watershed

(max. age)

Chestnut oak

White oak

Scarlet oak

Black oak

Mockemut hickory

Pitch pine

Shortleaf pine

Virginia pine

lake Winona Research Natural Area

Shortleaf pine

__ Carlson 1 99Sg

362

348

190

211

335

235

300

108

__ Stahle et al. 1985

__ Stable et al. 1985

Stable et al. 1985

108-314

Roaring Branch Research Natural Area

Shortleaf pine

Hot Springs National Park

Shortleaf pine

Number of IO-cm size classes h

John de la Howe tract

Shortleaf pine

Hardwoods

133-275

124247

This study

8

7

Scarlet Oak Natural Area Delapp and Wentworth d

Pitch pine

Hardwoods

Lake Winona Research Natural Area 1

2

Fountain 199 1

Roaring Branch Research Natural Area Fountain and Sweeney 1985

Shortleaf pine

Hardwoods

6

4

Kisatchie National Forest

Shortleaf and loblolly pines

Hardwoods

2 Martin and Smith 1991

7

5

D.b.h. of large trees (cm) ’

John de la Howe tract

Shortleaf pine

Loblolly pine

White oak

Hickory speciesg

Post oak

Southern red oak

Yellow-poplar

Roaring Branch Research Natural Area

Shortleaf pine

Standing coarse woody debris (no./ha)

John de la Howe tract

Snags 210 cm d.b.h.

Snags 250.8 cm d.b.h.

28 81

30 99

30 84

25 58

20 48

30 81

30 79

1 This study

Stahle et al. 1985

This study

36 66

31

5

170

40

21
Table 2 (metric units)-Standardized table of old-growth attributes for dry and dry-mesic oak-pine

forests (continued)

Quantifiable attribute Range4 Mean

Number

of standsb References

Standing coarse woody debris (no.ha) (continued)

Hot Springs National Park ’ 3 Johnson and Schnell 1985

(mass only-metric tons/ha)

Snags 27.6 cm d.b.h. 4.3 29.3

Downed coarse woody debris

John de la Howe tract

Volume (m’/ha)

Mass (metric tons/ha)

52

16.6

177

56.9

1 This study

Hot Springs National Park ’ 2.0 9.4

Tree canopy structure

John de la Howe tract

Number of canopy layers (l-3)

Percent canopy in gaps

Mean gap size’ (hectares)

2-3

24-80

.OOl-,204

37

,024

1 This study

a For the John de la Howe tract, this represents the range of means from areas showing different degrees of mortality, and applies to all attribute

data except age, diameter, and gap size.

bNumber of stands included (may not equal number of citations).

’ Oosting did not define the diameter limit for overstory. Density values are for overstory only. If understory is included, density of trees

22.5 cm in d.b.h. is 1697,220, and 380 per hectare for all trees, pines, and oaks, respectively. Basal area values are given for all trees X.5 cm

in d.b.h.

d Delapp, J.A., and T.R. Wentworth. 1977. Proposed research natural areas in the Southern Appalachian Mountains. 117 p. Unpublished

report. On file with: Highlands Biological Station, Highlands, NC 2874 1.

e Includes dominant and codominant trees that comprise the upper canopy. Mean diameter for trees from the John de la Howe tract is the

quadratic mean.

‘Includes mockemut, shagbark [Curya ovata (Mill.) K. Koch], and pignut hickory species.

g Carlson, P.J. 1995. An assessment of the old-growth forest resource on National Forest System lands in the Chattooga River Watershed.

Report to Chattooga Ecosystem Demonstration Management Project, U.S. Department of Agriculture, Forest Service, Region 8, Atlanta, GA.

83 p. On file with: USDA Forest Service, Southern Region, 1720 Peachtree Road, NW, Atlanta, GA 30367.

h For inclusion, 1 percent of all trees must bc present within a size class-size classes start at 10 cm.

i Average across three stands: (1) stand dominated by medium to large, actively growing shortleaf, (2) a “decadent” old-growth shortleafdominated stand, and (3) an old-growth oak-hickory stand with a few large shortleaf pines.

‘Refers to canopy gaps 1.4 ha in size. Gaps exceeding .4 ha were variable sized patches of extensive pine mortality that accounted for

58 percent of the total gap area.

22
Living Tree Component

Size structure-Density of trees 24 inches (10 cm) in d.b.h.

was similar for JDLH and two Ouachita Mountain sites but

varied greatly across the range of sites given in table 2.

Differences in density may be related to disturbance history

and age, as well as differences in diameter classes used to

describe the stands. Highest values were found in the Hot

Springs NP in Arkansas and lowest in the Kisatchie National

Forest of the Gulf Coastal Plain. The proportion of yellow

pines also varied greatly. The Lake Winona Research

Natural Area contained the highest density of pines and

comprised 90 percent of stand basal area and, therefore,

can’t be considered “oak-pine.” It is included here because it

is unique and includes dry and dry-mesic sites. Density of

large oaks and pines [220 inches (5 1 cm) in d.b.h.1 was

similar for the JDLH and Kisatchie National Forest and

exceeded values from two of the Ouachita Mountain sites.

Basal area was greater than 100 feet2 per acre (23 m2 per

hectare) for JDLH, Duke Forest, Scarlet Oak Natural Area,

and Lake Winona Research Natural Area (table 2). As with

density, basal area of large trees was almost identical for

JDLH and Kisatchie National Forest. The proportion of

basal area in species other than pine or oak was highest for

JDLH. Here, hickories accounted for 10 percent of the basal

area, followed by yellow-poplar, red maple, and sourwood.

The high end of the range for JDLH [ 150 feet* per acre (34

m* per hectare)] actually reflects stand-wide basal area

before significant pine mortality. This finding resulted when

recently dead pines were included in (live) basal area

calculations from an area of high pine mortality where

values exceeded 150. The high basal area may be due to

relatively mesic conditions at JDLH compared to typical dry

and dry-mesic sites, since just over half the JDLH was

classified intermediate, and the remainder, subxeric.

Maximum diameter-at-breast-height values of dominant and

codominant trees is limited to JDLH and Roaring Branch

Research Natural Area (table 2). Mean “large tree” diameter

at breast height for shortleaf was the same for both sites, but

Stahle et al. (1985) sampled trees to obtain long tree ring

chronologies, and, hence, these data may overestimate mean

diameter at breast height. The largest tree sampled at JDLH

was a 39-&h (99-cm) d.b.h. loblolly pine. The largest

shortleaf was 32 inches (8 1 cm) [shortleaf trees exceeding

32 inches (8 1 cm) were present in the stand but not in

sample plots]. The largest hardwoods were white oak [33

inches (84 cm)], southern red oak [32 inches (81 cm)], and

yellow-poplar [3 1 inches (79 cm)].

Detailed diameter-at-breast-height distributions were

available for JDLH and Lake Winona Research Natural

Area. Total stem distribution at Lake Winona Research

Natural Area was reverse-J (q = 1. l), typical of a balanced

uneven-aged system (Fountain 1991). However, distribution

of pines did not follow a normal or reverse-J distribution,

which Fountain suggested resulted from major disturbances.

Total stem diameter-at-breast-height distribution for JDLH

also followed a reverse-J curve (q = 1.6). Since the idealized

reverse-J distribution approximates a negative exponential

series, a straight line results when the logarithm of density is

plotted against diameter class (DeLiocourt 1898, Meyer

1952, Leak 1964). Implicit in this relationship is a constant

percentage loss with each increase in diameter. Diameter

distributions in many old or old-growth forests more closely

fit the form of the rotated sigmoid curve than the more

commonly expected negative exponential form (Goff and

West 1975, Schmalzer 1988). When plotted with log (10) of

density, the diameter-at-breast-height distribution of total

stems on JDLH approximates the rotated sigmoid curve (fig.

1). This pattern typically indicates high mortality for both

understory and oldest overstory trees, with little mortality in

actively growing canopy trees, which results in the flat

portion of the curve (Goff and West 1975). This curve may

shift to a straight line as more pines die (see “hardwoods

only” in figure 1).

Regardless of the specific form of the reverse-J distribution,

the single curve is the result of a combination of species

curves. Peak density of individual species curves is

expressed at different points along the diameter range

(Hough 1932), which, in turn, is determined by a number of

factors. Spatial and temporal variation in disturbance are two

of those factors. The result is patches of different size and

age, which support species with different light and space

requirements.

Hardwood-dominated forests, examples of the uneven-aged,

“steady state” phase, have been described by Peet and

Christensen (1987) as a mosaic of even-aged patches of

various sizes and ages. Within each gap or patch, the early

phases of forest development (establishment, thinning, and

transition) take place. This process of maintaining the

composition and uneven-aged condition through time is

applicable to oak-pine types only if sufficient canopy and

forest floor disturbance occurs periodically to establish

patches of oak and pine species. The development stage

exemplified by JDLH could be described as the transition

phase (Peet and Christensen 1987) or “understory

reinitiation” phase (Oliver 198 l), which precedes the “old

growth” phase, respectively. An alternative view is that the

JDLH stand, where pines and hardwoods have persisted for

200 years, represents an ephemeral old-growth stage. The

change in composition and structure of the JDLH stand

discussed above and shown in figure 1 illustrates the

temporal nature of old-growth oak-pine. Although it may be

temporal on a given site, it is maintained on the broader

landscape by disturbances distributed over space and time.

Age structur+Age ranges or maximum ages, or both, are

presented for several areas in table 2. In the Great Smoky

23
Diameter distribution - John de la Howe Forest

n All species

o Hardwoods only

15 20 25 30

Diameter class (inches)

Figure l-Diameter distribution for hardwoods and all species (hardwood and pines). Data is from the John de la Howe

Tract, McCormick County, South Carolina. Stems per acre (log 10 transformation) versus diameter [2-inch (S-cm)

diameter classes]

Mountains NP, white and chestnut oaks and pignut hickory

were the oldest oak-pine species exceeding 300 years,

whereas pitch and Table Mountain pines exceeded 200

years. Shortleaf pine exceeded 300 years in the Ouachita

Mountains (Stahle et al. 1985) and shortleaf, white oak, post

oak, and hickories exceeded 200 years at JDLH. It is

noteworthy that shortleaf and several oaks (especially white

oak) have been observed to respond to crown release at 180

years old and that the maximum age of black and scarlet

oaks in the Southern Appalachians exceeds that commonly

observed for these species (see footnote 7).

The uneven-aged structure of stands at JDLH and the

Ouachita Mountains, described in the previous section, is

also supported by age class distributions (Stahle et al. 1985).

Three Ouachita Mountain old-growth forests contained

shortleaf pine in several age classes of 100 years or greater.

Similar observations were made for this species at JDLH

(fig. 2). White and post oaks, shortleaf and loblolly pines,

and hickories were found in most age classes, and there were

also a few old southern red oak and yellow-poplar.

Loblolly was more common than shortleaf in age classes

~150 years at JDLH. Two distinct groups of white oak were

observed. The youngest group ranged from 75 to 100 years

and varied in size from 5 to 20 inches (12.7 to 50.8 cm) in

d.b.h., whereas the older group ranged from 140 to 200

years and 17 to 33 inches (43.2 to 83.8 cm) in d.b.h. Most

other species, including red maple, red oaks, and most

yellow-poplar sampled, were generally less than 100 years

old. With the exception of pines, very few individuals were

in the lOO- to 140-age class. The presence of more mesic

and fire-sensitive species like red maple I 100 years in age

suggests that fire may have affected this stand before the late

1800’s. Age distributions of other species suggest scattered

disturbance of sufficient size to result in regeneration of pine

and other intolerant species over the past 200 years.

Canopy structure-In most old-growth oak-pine forests,

there are two to three distinct canopy layers, in addition to

an herbaceous layer (table 2). Typically, there is an

ericaceous shrub layer dominated by mountain laurel,

bluebetry, or hucklebeny; a midstory layer with variable

amounts of blackgum, dogwood, sourwood, Florida maple,

and eastern hophornbeam; and the dominant canopy layer.

24
John de la Howe Forest: age distribution by species group

I3-

I-

)---

Loblolly

&j Shodleaf

[7 White oak

Hickory spp.

I Red oaks I Red maple

I Yellow-poplar m Other spp.

1

50-74 75-9s loo-124 125-149 150-174 175+

25year age class

Figure 2-Age class distribution for major species from the John de la Howe Tract, McCormick

County, South Carolina.

25
Height information is lacking. The mean height of dominant

hardwoods and pines in the JDLH stand was 100 and 110

feet (30.5 and 34.4 m), respectively. The mean height of

midstory species (hardwoods only) was 48 feet (14.6 m).

The understory structure and composition in old-growth

oak-pine have likely been impacted by 20’-century fire

suppression as evidenced by a greater presence of mesic

species (Tucker 1991).

The age structure of trees in a forest is the sum of the age

structures in the mosaic of patches of the forest, and

descriptions of the mosaic may provide more information

about the dynamics of the system than an overall forest

assessment (Harper 1977). Information on canopy gaps is

available only for JDLH (table 2). Mortality patterns or

distribution of canopy gaps has been described previously

by White and Lloyd (1995). They collected data at four

levels of mortality defined as canopy openness or percent of

canopy in gaps: (1) low, ~25 percent canopy gaps; (2)

intermediate, 25 to 50 percent canopy gaps; (3) high-old,

>50 percent canopy gaps formed 3 or more years prior; and

(4) high-recent, >50 percent in canopy gaps occurring within

the past 1 to 2 years. In the areas of low and medium

mortality, the canopy is characterized as relatively

continuous, broken by various sized canopy gaps, and

generally ~0.3 acres (.12 ha), averaging 0.06 per acre (.024

per hectare) [58 feet (17.7 m) in diameter]. Most mortality is

in pine; 23 percent is in hardwoods, and 15 percent is in

pines and hardwoods combined. The range and mean gap

size found at JDLH on low-to-medium mortality areas was

similar to that found by Clinton et al. (1993) in a mature

Southern Appalachian mixed oak forest. In the highmortality areas (recent and old), overstory canopy openness

ranged from 60 to 80 percent, with open patches from 0.5 to

10 acres (.2 to 4.0 ha). Almost all mortality was due to

shortleaf and loblolly pines. Because pine mortality occurred

at different times in the life of the stand and because variable

amounts of hardwoods were present during pine mortality,

the size and age structure within gaps varies across the

stand.

Dead Tree Component: Coarse Woody Debris

Across the gradient of canopy openness (mortality)

described for JDLH, certain old-growth characteristics vary

greatly. Density of standing coarse woody debris (CWD) or

snags 24 inches (10 cm) in d.b.h. ranged from about 15 per

acre (37 per hectare) in the areas of low-to-medium

mortality to 70 per acre (170 per hectare) in the areas of

high-recent mortality (table 2). In Kentucky, McComb and

Muller (1983) reported 14 to 32 snags [24 inches (10 cm) in

d.b.h.1 per acre (35 to 79 per hectare) in old-growth beech

(Fugus grandijblia Ehrh.) and chestnut oak forests,

respectively. Sabin (199 1) reported 11 snags per acre (27

per hectare) across a range of 20- to 60-year-old South

Carolina Piedmont forest types. The density of large snags

[220 inches (50.8 cm) in d.b.h.1 averaged 8 per acre (20 per

hectare) at JDLH, exceeding that described for old-growth

and second-growth mixed mesophytic forests in Kentucky

@&Comb and Muller 1983). The only other snag data for

old-growth oak-pine is in terms of mass rather than density

(Hot Springs NP) (Johnson and Schnell 1985) and,

therefore, are not comparable to those reported for JDLH.

The pattern for JDLH snags also applies to downed CWD,

except that the high-recent mortality area currently has

relatively low quantities. In the high-recent mortality area,

most of the dead pine is still standing in contrast to high-old

mortality areas. Values of 25 tons per acre (57 metric tons

per hectare) in the high-old mortality class are higher than

the 8 to 11 tons per acre (18 to 25 metric tons per hectare) in

old-growth mixed oak-hardwood forests of the Eastern

United States (Lang and Forman 1978, MacMillan 198 1,

Harmon et al. 1986, Muller and Liu 1991), but less than half

of that reported for mature uneven-aged pine on the South

Carolina Coastal Plain following Hurricane Hugo (Myers et

al. 1993). Coarse woody debris mass for Hot Springs NP

(Johnson and Schnell 1985) is much smaller than that for

JDLH. Inputs of CWD from broad-scale disturbances, such

as insect-related mortality and hurricanes, represent large but

infrequent pulsed inputs.

Coarse woody debris in the low-to-medium mortality areas

correspond to the range for eastern old-growth oak forests

listed above. This information, coupled with gap information

described earlier, implies that disturbance patterns and

associated effects that predominate in low-to-medium

mortality areas are similar to those found in other mature or

old-growth forests in the East. This is temporary, however,

given that additional pine mortality is expected in these

areas in the future. As pines succumb to southern pine

beetle, littleleaf disease, and pathogens, a further shift to

oaks and other hardwoods will occur. Without forest floor

disturbance such as fire, pines will become minor

components of this stand in the future.

In the Southern Appalachians, snags, downed CWD, and

other old-growth characteristics, such as pit and mound

microtopography, may be reduced or lacking in the drier,

upland old-growth types (see footnote 7). The deviation

from the classic old-growth characteristics attributed to cove

hardwood stands is most evident on xeric and subxeric sites

where disturbances (especially fire) are more frequent and

intense.

26
Forest Dynamics and Ecosystem Function

Presettlement Disturbance Regimes

A review of paleoecological and other literature pertinent to

the history of the oak-pine forest type (White and Lloyd

1995) revealed three important points: (1) oaks and pines

have been components of the eastern deciduous forest for

thousands of years, (2) shifts in their relative abundance

have been related to climate changes and to fire used by

aboriginal people in the past 10,000 years, and (3) oaks and

pines that dominate the dry and dry-mesic oak-pine type

require distirbance to maintain their dominance. Buckner

(1989) described the pine-hardwood type as a “mid-seral

stage that is ephemeral on a given site and is maintained in a

changing landscape mosaic where scattered disturbances

reinitiate succession in a stochastic manner.” In this paper,

we have referred to the old-growth oak-pine type as

resembling the transition or understory reinitiation phase of

forest development or the mid-to-late phase of plant

succession. It is, indeed, ephemeral on a given site unless

sufficient disturbance occurs. It is important to

acknowledge, however, that multiple ecological outcomes

are possible for comparable sites (Oliver 198 1, Sprugel

1991). The “outcome” depends on the nature of the

disturbance and its interaction with the biotic and abiotic

components of a given site.

Disturbance continuum-A greater understanding of

presettlement disturbance and species response to

disturbance should clarify management options for

recognizing, creating, and maintaining old-growth types on

the landscape. Our understanding of these factors, however,

is limited. Natural forest disturbance in the form of

hurricanes, tornadoes, ice and wind storms, drought, insectand pathogen-induced mortality, and lightning fire represent

the low end of a disturbance continuum. The presettlement

human disturbance regime superimposed on the natural

disturbance regime represents the upper end of this

continuum in which eastern forests have developed.

Although presettlement disturbance dynamics are much

more complex than the above description implies,

distinguishing endpoints of a disturbance continuum may

represent a starting point for improving our understanding.

With increased urbanization in the Eastern United States,

accompanied by increased demands on Federal land,

management options are more limited (e.g., restrictions on

burning to minimize smoke). However, if we know more

about the endpoints of this disturbance continuum,

managers, decision-makers, and the public will be in a better

position to make decisions as to the assemblage of

vegetation types we seek to manage or restore.

The relative impact of nonhuman and human disturbances

varies across regions. Lightning fires are more prevalent in

the Coastal Plain and Ozark region (Komarek 1964, 1968).

Hurricanes are more common in the Coastal Plain than in

inland areas, whereas tornadoes are common events in the

Ouachitas and Ozarks (Turner 1935, Foti and Glenn 1991,

Tucker 199 1). The impact of early human disturbance was

likely greater in regions where the frequency and impact of

lightning fires were less such as the mountains and Piedmont

(Frost 1995). Earlier, we described pyrogenic patches in

parts of the Coastal Plain tinctioning as ignition sources for

adjacent communities. Although this situation may apply to

other regions (e.g., mountainous regions:xeric ridgetops), it

may be less relevant for the Piedmont where moisture

gradients are less extreme than in areas with greater

topographic relief or areas with more coarse textured soils,

or both. Compared to other regions, we have little

understanding of the spatial and temporal aspects of

presettlement disturbance in the Piedmont.

Fire frequency-Various estimates of fire frequency have

been given for different physiographic regions earlier in this

paper, but the variability associated with these estimates is

often large (Christensen 1988b). Christensen suggests that

this variability is an essential component in maintaining fireprone ecosystems by citing the importance of chance

variations in determining structure, composition, and

distribution of different ecosystems. In the Ouachitas, firereturn intervals varied from 7.25 to 32 years depending on

site and period of time examined (Johnson and Schnell

1985, Foti and Glenn 199 1). Fire-return intervals were

estimated to be 5 to 15 years for shortleaf pine-oak-hickory

of the Louisiana Coastal Plain (Martin and Smith 199 1).

Frost (1995) describes presettlement fire frequency for the

Southeastern United States as being 1 to 3 years for the

southern part of the Atlantic Coastal Plain and lower Gulf

Coastal Plain; 4 to 6 years for the Coastal Plain of Virginia

and upper North Carolina, the Piedmont, and the middle and

upper parts of the Gulf Coastal Plain; 7 to 12 years in

northern Alabama and part of the Appalachians, foothills,

and upper Piedmont [300 to 3,000 feet (92 to 915 m) in

elevation]; and 12 or more in the high mountains (6,000 feet

or 1829 m) and wet swamps. Nowacki (1995) estimated the

presettlement disturbance frequency (includes fire, storms,

drought, disease) in the Valley and Ridge province of

Pennsylvania to be 2 1 years.

Patch size-Successful establishment of pines, and to a

lesser degree oaks, depends on ground fires (Christensen

198 1, Abrams 1992, Van Lear and Watt 1993).

Establishment of these species as canopy dominants,

however, depends on the amount of canopy disturbance. In

the Gulf Coastal Plain, small-to-intermediate-sized gaps are

thought to be required for maintaining oak-pine and related

types (Harcombe and Marks 1978, Golden 1979, Martin and

Smith 199 1). In the Ouachitas, large-scale disturbances were

found to be the origin of three “virgin” shortleaf pine stands

(Turner 1935). Turner suggested tornadoes were

responsible, but it could have been a combination of other

27
canopy disturbance and fire as well. Carlson (see footnote 7)

described canopy-age classes exceeding 150 years across the

Southern Appalachians, suggesting region-wide, prelogging

disturbances. These were common on upper slopes and

ridges currently occupied by oaks or pines, or both, and

were assessed as relatively large-scale disturbances.” At

JDLH, a range of canopy gap sizes resulted from mostly

pine mortality, but pine regeneration was considered

unlikely without fire or other forest floor disturbance (White

and Lloyd 1995). If pine regeneration is present, canopy

disturbances resulting in 0.1 acre (.04 ha) or larger openings

[74 feet (22.6 m) in diameter] are sufficient for

establishment of yellow pines in the Piedmont (Waldrop

199 1, Perry and Waldrop 1995). Beyond a certain minimum

gap size, fire is crucial for oak-pine maintenance.

Consequently, a wide range of disturbance patch sizes may

result in pine establishment if forest floor disturbance is ,

sufficient.

Ecological Processes

Early concepts of old-growth forests were based on the

steady-state, equilibrium view of forest dynamics, but

several authors have presented alternative views of forest

succession or have acknowledged the role of a wide range of

disturbances in-forest development (Egler 1954, Spurr and

Barnes 1973, Harper 1977, Oliver 1981, Denslow 1985,

Pickett and White 1985, Glitzenstein et al. 1986,

Christensen 1988b, Platt and Schwartz 1990, Sprugel 1991).

The dry and dry-mesic oak-pine type and some other oldgrowth types do not adhere to steady-state dynamics; rather,

disturbance by fire or other agents is required for its

maintenance on the landscape. Various sized patches of

various ages need to be reestablished on the landscape to

allow forests to continually recruit into oak-pine old growth.

If not, oak-pine old growth will be lost over much of its

former range.

Wind, fire, insects, and plant pathogens commonly impact

ecological processes in old-growth oak-pine forests, and

they often interact adding to ecosystem complexity (Jackson

1968). Broad-scale wind and insect disturbance can produce

large, pulsed inputs of CWD. If ignition occurs when fuel

loads are high and conditions are conducive to fire, severe

fires can occur. This can significantly impact future

composition of the forest and affect both short- and longterm quantities of nutrients and organic matter on the site

even though tires rarely consume all CWD. In the absence of

fire, decay proceeds, rendering CWD less conducive to

supporting tire. This large-pulsed input of CWD may

represent habitat for many organisms and contributes much

organic matter to the site. Coarse woody debris retains water

” Personal communication. 1995. Paul J. Carlson, Forester, 162 Goshen

Road, Franklin, NC 28734.

and serves as both nutrient source and sink during the

decomposition process, playing a critical role in water and

nutrient cycles. With relatively frequent ground fires, CWD

may have been a much less significant component of oldgrowth dry and dry-mesic oak-pine during presettlement

times.

The dynamics of CWD inputs and the long-term effects on

various ecosystem processes are not well understood (Van

Lear 1996, Waldrop 1996). We have provided an example

of large-pulsed inputs of CWD due to stand-wide pine

mortality. In simulation studies of CWD dynamics for mixed

hardwood stands growing on xeric and mesic sited in

Tennessee, CWD loads increased from age 32 to 90

followed by a gradual decrease through age 200 (Waldrop

1996). Waldrop demonstrated that whereas CWD inputs

were much higher on the mesic site than on the xeric site,

CWD loads were similar when a slightly slower

decomposition rate was simulated for the xeric site. Inputs

from natural disturbance, such as insect-related pine

mortality, was not included in the simulation but might have

a significant impact on long-term CWD dynamics.

If a management objective is to maintain an old-growth

forest type that contains a significant pine component, it is

important to consider the effect of insects and pathogens on

the composition and structure of old growth (Haack and

Byler 1993), as well as the interaction of old growth with

surrounding stands. In the Southeastern United States, the

impact from southern pine beetle is less in mixed pinehardwood stands than in pure pine stands because pine

continuity is interrupted by hardwoods (Belanger et al. 1986,

Showalter and Turchin 1993). The management challenge is

to allow natural disturbance in old-growth forests, while

minimizing losses in surrounding stands. Separating

intensively managed stands (high pine component, disease

risk dependent on stand conditions) from old-growth oakpine stands (moderate pine component, moderate-to-high

disease risk) with pure hardwood stands or with mixed pinehardwood stands may reduce disease-caused losses (White

and Lloyd 1995).

Postsettlement changes-The most dramatic postsettlement

change to eastern landscapes was conversion of forests to

agricultural fields, especially widespread in the Piedmont

and Coastal Plain. Few areas were untouched by the plow,

and soil erosion was extensive in some areas, resulting in

significant changes to site productivity and biological

potential. Other major postsettlement changes included the

shift to woodland grazing by cattle and hogs at the time of

settlement (shift from native grazers such as bison and elk),

the cessation of woodland grazing by 1900, and 20’-century

fire suppression. These changes likely contribute to

differences in composition and structure between

presettlement and current old-growth forests (White and

White 1996). It is important to acknowledge the impact of

28
these disturbances on the distribution and composition of

current and future old-growth forests.

Without fire, species less tolerant of fire, such as red maple,

Florida maple, and eastern hophombeam, increase in

abundance. Loblolly pine and other mesic species have been

described as “creeping” upslope due to lack of fire. This

may also apply to white pine in the mountains. Plant

diversity in most dry and dry-mesic oak-pine forests is

relatively low, especially since many of these types have

understories dominated by one or two layers of ericaceous

species. Understory diversity in presettlement oak-pine types

may have been greater than current stands (Harper 1943,

Frost 1993). Frost described presettlement oak-pine types of

the upper Coastal Plain as woodland fire types with a

species-rich herb layer. Harper described the shortleafpineoak forests of the dry uplands of Alabama as having

lightning fires every 10 years and having a “good deal of

grassy undergrowth,” suggesting a potentially more diverse

herbaceous layer than is currently seen. How prevalent this

condition was in other regions is unknown. Logging

disturbance may result in oak- and pine-dominated forests,

but will not create the diverse, pyrogenic understories Frost

refers to.

When Does Old Growth Begin?

Defining old growth demands the following questions be

addressed: (1) when is a forest considered old growth, and

(2) what is the minimum area needed to maintain old-growth

forests? The first question is partially clarified by the generic

definition of old growth, as drafted by the National OldGrowth Task Group in 1989:

Old-growth forests are ecosystems distinguished by

old trees and related structural attributes. Old-growth

encompasses the later stages of stand development

that typically differ from earlier stages in a variety of

characteristics which may include tree size,

accumulations of large dead woody material, number

of canopy layers, species composition, and ecosystem

function.

No single characteristic, such as age or size, can define old

growth. To illustrate this, consider a 125- to 150-year-old

oak-shortleaf pine stand that contains many of the structural

and compositional features of old growth but may not

exhibit accumulation of dead woody material. The first

significant CWD accumulation in this stand could have

resulted from a severe southern pine beetle/littleleaf

epidemic that essentially eliminated the pine component.

How does this disturbance impact the old-growth status of

this stand? Can it be defined as old growth because snags

and downed CWD are “suddenly” present in the stand, or is

it no longer old growth because it lost the old pine

component? We might consider that it was old growth

before and after the disturbance, depending on other stand

factors, and that we have witnessed an abrupt compositional

change in the old-growth stand from oak-pine to oak.

We consider oak-pine stands that have minimal evidence of

postsettlement human disturbance and contain pines

exceeding 100 to 125 years to be approaching, if not already

functioning as, old growth. For Virginia pine on most sites,

100 years represents old age. For shortleaf pine on good

sites, this may be too young to be considered old growth.

Hepting (1971) specifies 80 years for maturity of Virginia

pine, 100 years for loblolly and pitch pines, and 120 years

for shortleaf pine. In general, we recommend that most

stands with pines and oaks that exceed 100 to 125 years and

have experienced little recent human disturbance, be

considered to be in the early stages of old growth. Impacts

from disturbance, site quality, growth form of the oldest

cohort, age and size structure, composition, and

accumulation of dead woody material must all be considered

when characterizing the old-growth status of forests.

Addressing the second question regarding the area required

to maintain old growth is more complex. When large-scale

disturbances are associated with a forest type, the area of

land required to maintain that type on the landscape

increases accordingly (Shugart 1984, Busing and White

1993). Also, the probability that a stand will be subjected to

natural catastrophes increases with stand age (Oliver 198 1).

Rather than focusing on the size of a stand, the entire

landscape should be emphasized, especially with ephemeral

forest types. Whether an old-growth oak-pine stand is 5 or

100 acres (2 or 40 ha), it matters little if it is the only oldgrowth patch within some larger area [e.g., 5,000 to 20,000

acres (2000 to 9000 ha)]. To ensure that old-growth forests

are maintained on the landscape, a mosaic of relatively

large, contiguous blocks of old-growth forests among blocks

of younger forests are needed. Managing for this mosaic

better ensures that the structural, functional, and

compositional diversity of forested ecosystems will be

maintained (Harris 1984, Franklin and Forman 1987,

Franklin 1988). Since many public land managers manage

forests for multiple values, strategies to maintain old-growth

forests and to produce other sustainable commodities should

be integrated.

Information Gaps

Gaps in our knowledge about old-growth oak-pine result

from the fact that few of these stands exist and even fewer

have been studied. There is a need to locate, document, and

describe existing stands and their old-growth characteristics.

In addition, more work is needed to document threatened,

endangered, and sensitive plant and animal species that are

associated (currently or historically) with this forest type.

Information is also lacking on presettlement forest

conditions, spatial and temporal scales of nonhuman

disturbances, and the distribution of Native American

29
populations and their impact on presettlement landscapes.

The availability of information varies with the region. In

some cases, information on Native American populations

and activities may be well known, but this information has

not been put into a larger spatial and temporal context.

Representative Old-Growth Forests

Dry and dry-mesic old-growth oak-pine forests verified

directly or indirectly by the authors

Hot Springs National Park, Arkansas

Lake Winona Scenic Area, Ouachita National Forest,

Arkansas

Roaring Branch Research Natural Area, Ouachita

National Forest, Arkansas

Bobs Creek Shortleaf and Old Shortleaf Slope on the

Kisatchie National Forest, Louisiana

Great Smoky Mountains National Park, North Carolina

and Tennessee

Linville Gorge, Pisgah National Forest, North Carolina

Scarlet Oak Natural Area, Pisgah National Forest, North

Carolina

John de la Howe Tract, McCormick County, South

Carolina

Other dry and dry-mesic old-growth oak-pine forests not

verified by the authors

.

.

.

.

.

a

.

.

.

.

0

Magazine Mountain, Logan County, Arkansas

Marshall Forest Preserve, west of Rome, Georgia

Ack Tract, along Piney River, Texas County, Missouri

Big Spring Towering Pines, Carter County, Missouri

Hickory Canyons Natural Area, Ste. Genevieve County,

Missouri

Meramec Upland Forest Natural Area, Meramec State

Park, Missouri

Mudlick Mountain Natural Area, Sam A. Baker State

Park, Missouri

Bienville Pines Scenic Area, Bienville National Forest,

Mississippi

McCurtain County Wilderness Area, McCurtain County,

Oklahoma

Martins Hill, Buchanan State Forest, Bedford County,

Pennsylvania

Lennox Woods Preserve, Red River County, Texas

Acknowledgment

We would like to thank John Haney, Carol Perry, Shawna

Reid, and Jane Thompson for field assistance at John de la

Howe. We would also like to thank the John de la Howe

staff for their cooperation. Critical reviews of the manuscript

were provided by Peter Kapeluck, Greg Nowacki, Jane

Thompson, Joan Walker, Tom Waldrop, and Peter White.

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34
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35
Appendix A

List of common trees, shrubs, and herbaceous species” occurring in old-growth and non-old-growth forest stands representative of the oak-pine type in

the Eastern United States, with scientific and common names given. Nomenclature is from Kartesz and Kartesz (1980).

I

Old-growth stand$ Non-old-growth standsb

Piedmont Gulf CP Ouachita Mountains Southern Appalachian Mountains Plateau

Common name Scientific name

I 2 3 4 5 6 7 8 9

JDLH Duke KN!: RBRNA HSNP LWRNA SCONA ESCARP Chauga S::PP S&MP Sh%-H

14

Pine Mt.

-

Trees: others (cont.)

Winged elm

Southern sugar maple

Persimmon

Yellow-poplar

Black cherry

Sassafras

American elm

American chestnut

Black locust

Shrubs: Ericaceae

Blueberry

Deerbeny

Lowbush blueberry

Sparkleberry

Huckleberry

Danglebeny

Mountain laurel

Wild azalea

Wild azalea

Highbush blueberry

Trailing arbutus

Wintergreen

Bear huckleberry

Male blueberty

Fetterbush

Ulmus alata Michx.

A. jloridanum (Chapman)

Pax

Diospynrs virginiana L.

Liriodendron tulipifera L.

Prunus serotina Ehrh.

Sassafras albidum

(Nutt.) Nees

U. americana L.

Castanea dentata

(Marsh.) Borkh.

Robiniapseudoacacia L.

Vaccinium spp.

V. stamineum L.

V. vacillans Torr.

V. arboreum Marsh.

Gaylussacia baccata (Wang.)

K. Koch

G.fiondosa (L.) T.&G.

Kalmia latifolia L.

Rhododendron canescens

(Michx.) Sweet

R. periclymenoides

(Michx.) Shinners

Y. elliottii Chapman

Epigaea repens L.

Gaultheriaprocumbens L.

Gaylussacia ursina (M.A.

Curtis) T.&G.

Lyonia ligustrina (L.) DC.

Pieris Joribunda

(Pursh) B. & H.

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

(continued)

I

Appendix A

List of common trees, shrubs, and herbaceous species” occurring in old-growth and non-old-growth forest stands representative of the oak-pine type in

the Eastern United States, with scientific and common names given. Nomenclature is from Kartesz and Kartesz (1980).

I

Old-growth stands* Non-old-growth stand&

Piedmont Gulf CP Ouachita Mountains Southern Appalachian Mountains Plateau

1 2 3 4 5 6 7 8 9 IO II

SM:VP SM:PP ShblMP SIv&-H

14

Common name Scientific name JDLH Duke KNF RBRNA HSNP LWRNA SCONA ESCARF’ Chauga Pine Mt.

VilleS

SlllilaX

Virginia creeper

Grape

Muscadine

Yellow jessamine

Trumpet honeysuckle

Sawbrier

Greenbrier

r;:

Herbs: Asteraceae

sunfl0we.r

PUSSytOeS

Aster

Hawkweed

Hawkweed

Goldenrod

Anomalous aster

Elephant’s_foot

Daisy fleabane

Prairie sunflower

Stiff sunflower

Blazing star

Ciumweed

Goldenrod

Hispid goldenrod

Ironweed

Tickseed

Smiler spp.

Parthenocissus quinquefolia

(L.) Planch.

Vitis spp.

V. rotundifolia Michx.

Gelsemium sempervirens

(L.) St. Hill.

Lonicera sempervirens L.

S. glauca Walt.

S. rotundifolia L.

Helianthus spp.

Antennaria plantaginijolia

L. Richards.

Aster spp.

Hieracium gronovil L.

H. venosum L.

Solidago spp.

A. anomalus Engehn.

Elephantopus tomentosus L.

Erigeron strigosus

Muhl. ex Willd.

H. laetifronrs Pers.

H. divaricatus L.

Liatris spp.

Silphium spp.

Solidago caesia L.

S. hispida Muhl.

Vernonia spp.

Coreopsis major Walt.

(continued)
Appendix A

List of common trees, shrubs, and herbaceous species’ occurring in old-growth and non-old-growth forest stands representative of the oak-pine type in

the Eastern United States, with scientific and common names given. Nomenclature is from Kartesz and Kartesz (1980).

Old-growth stands”

!

Non-old-growth stand&

Piedmont Gulf CP Ouachita Mountains Southern Appalachian Mountains Plateau

Common name Scientific name

I 2 3 4 5 6 7 8 9

JDLH Duke KNF RBRNA HSNP LWRNA SCONA ESCARP Chauga SG:PP S&VIP

Herbs: Asteruceae (cont.)

Golden aster

Goldemod

Goldenrod

Herbs: Fabaceae

Beggar%-lice

Tephrosia

Baptista

Partridge pea

Butterfly-pea

Beggar’slice

Lespedeza

Wild indigo

Butterfly-pea

Dollar leaf

Herbs: grasses and sedges

All grass species

Panic-grass

Broom-straw

Broom-sedge

Sedge

Triple-awned grass

Poverty-grass

Little bluestem

Nutrush

Indian grass

Pityopsb graminifolia

(Michx.) Nutt.

S. bicolor L.

S. oroda Ait.

Desmodium spp.

Tephrosia spp.

Baptisia spp.

Cassiafasciculata Michx.

Centrosema virginianum

(L.) Benth

D. laevigafum (Nutt.) DC.

Lespedeza spp.

B. tintoria (L.) R. Br.

Clitoris mariana L.

D. rotundifolium DC.

Panicurn spp.

Andropogon spp.

A. virginicus L.

Carex spp.

Aristida spp.

Danthonium spp.

D. spicata (L.) Beauv.

ex. Roemer & Schul.

Paspalum spp.

Schizachyrium scoparium

(Michs.) Nash

Scleria spp.

Uniola spp.

Sorghastnrm nutans

(L.) Nash

X

X

X

X

X

X

X

X

X

XC

c

s

XC

XC

X

X

X

XC

x”

X

X

X

X

X

X

X

X

Sh&-H

14

Pine Mt.

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

(continued)
Appendix A

List of common trees, shrubs, and herbaceous species” occurring in old-growth and non-old-growth forest stands representative of the oak-pine type in the

Eastern United States, with scientific and common names given. Nomenclature is fromJCartesz and Kartesz (1980).

Common name

Herbs: others

Pipsissewa

Bracken fern

Dwarf Dutchman’s-pipe

Wild yam

Bedstmw

Wild ginger

Milkweed

Prairie spurge

Wild ginger

Partridge-berry

Solomon’s-seal

Five-fmger

False Solomon%-seal

Purple cress

Perfoliate bellwort

Colicroot

False foxglove

Bellflower

Galax

Gentian

Scientific name

Old-growth standsb Non-old-growth standab

Piedmont Gulf CP Ouachita Mountains Southern Appalachian Mountains Plateau

I 2 3 4 5 6 7 8 9 14

JDLH Duke KNF RBRNA HSNP LWRNA SCONA ESCARP Chauga Shk’P &:PP Sh&MP &2-H Pine Mt.

-

Chimaphila maculata

(L.) Pursh

Pteridium aquillinum

(L.) Kuhn

Aristolochia serpen taria L.

Dioscorea villosa L.

Galium spp.

Hexastylus spp.

Asclepias spp.

Euphorbia corollata L.

H. virginica (L.) Small

Mitchella repens L.

Polygonatum biflorum

(Walt.) Eli.

Potentilla canadensis L.

Smilacena racemosa

(L.) Deaf.

Streptanthus maculatus

Nutt.

Uvulariaperfoliata L.

Aletris farinosa L.

Aureolaria laevigata

(Raf.) Raf.

Campanula divaricata

Michx.

Galax urceokata (Pair.)

BlUNIIitt

Gentiana villosa L.

X

X

X

X

X

X X

X

x -

x -

X X

X

(continued)

.__....._ I
Appendix A

List of common trees, shrubs, and herbaceous species’ occurring in old-growth and non-old-growth forest stands representative of the oak-pine type in

the Eastern United States, with scientific and common names given. Nomenclature is from Kartesz and Kartesz (1980).

Common name Scientific name

Old-growth stands” Non-old-growth stands*

Piedmont Gulf CP Ouachita Mountains Southern Appalachian Mountains Platl%W

I 2 3 4 5 6 7 8 9 14

JDLH Duke KNF RBRNA HSNP LWRNA SCONA ESCARP Chauga ST&P &:PP Sh&MP Si&-H Pine Mt.

Herbs: others (cont.)

Stargrass Hypoxir hirsuta (L.)Coville - - - - - - - - - - - - - X

Violet Viola spp. X

Birdfoot-Violet V. pedata L. X_ --- _ X

Violet V. sagittala Ait. X

a Species are listed by species group in the order of their abundance across old-growth stands.

* Stands 1 through 14 ate: JDLH = John de la Howe, mixed oak-shortleaf/loblolly pine forest (this study); Duke = Duke Forest, white oak-post oak-shortleaf pine (Oosting 1942); KNF = Kisatchie

National Forest, shortleaf pine-oak-hickory forest (Martin and Smith 1991); RBRNA = Roaring Branch ResearchNatural Area, oak-shortleaf pine forest (Fountain and Sweeney 1985); HSNP = Hot

Springs National Park, oak-hickory-shottleaf pine (Dale and Watts 1980); LWRNA = Lake Winona Research Natural Area, shortleaf pine forest (Fountain and Sweeney 1987, Fountain 1991);

P

SCONA = Scarlet Oak Natural Area, scarlet oak-pitch pine forest (Delapp and Wentworth 1977); ESCARP = Blue Ridge Escarpment, Thompson River Gorge, pitch pine-scarlet oak/oak-pitch pine

(Racine 1966); Chauga = Chauga River Gorge, shottleaf pine forest (Tobe et al. 1992); SM:VP = Great Smoky Mountains National Park, Virginia pine forest (Whittaker 1956); SM:PP = Great Smoky

Mountains National Park, pitch pine forest (Whittaker 1956); SM:TMP = Great Smoky Mountains National Park, Table Mountain pine forest (Wbittaker 1956); SM:C-H = Great Smoky Mountains

National Park, chestnut oak-chestnut-heath forest (Whittaker 1956); Pine Mt. = Pine Mountain, Kentucky, pine-oak to oak-pine forests (Braun 1935).

’ Species not sampled in old-growth stands within the Kisatcbie National Forest (KNF) but were described as part of the KNF shortleaf pine oak-hickory natural community type by Martin and Smith

(1991).
Appendix B

Descriptions of old-growth stands and data used in table 2 by physiographic region:

Piedmont

(1) John de la Howe tract, a mixed oak-shortleaf-loblolly stand near McCormick, SC, has

been described in part by White and Lloyd (1995) (see text for additional information);

(2) a white oak-post oak stand on the Duke Forest, NC (Oosting 1942). This stand, which

contained a significant shortleafpine component, was one of three that Oosting sampled

to characterize the white oak-post oak type that he considered a variant of the oak-hickory

climax. Oosting described the oak-hickory stands as having 200- to 300-year-old trees

and showing little evidence of recent disturbance.

Ouachita Mountains

Three stands from this region were described previously as old growth in the literature:

(1) Roaring Branch Research Natural Area (Fountain and Sweeney 1985), (2) Lake

Winona Research Natural Area (Fountain 1991), and (3) Hot Springs National Parkcomposition and structure (Dale and Watts 1980) and coarse woody debris (Johnson and

Schnell 1985). Age data were derived from Stahle et al. (1985) for these areas.

Gulf Coastal Plain

Shortleaf pine-oak-hickory stands in the Kisatchie National Forest, LA, were described as

old growth by Martin and Smith (199 1). The two stands used in table 2 are referred to as

Bobs Creek Shortleaf and Old Shortleaf Slope. They are 113 and 127 years old,

respectively.

Southern Appalachians

Two stands in the Scarlet Oak Natural Area were characterized in the disturbance class as

“virgin, undisturbed” (see footnote 8). The report contained composition and structure

data for each stand. We relocated one of these stands and obtained increment cores from

chestnut oak, scarlet oak, and pitch pine. Pitch pine and chestnut oak exceeded 150 years

old, but younger age classes of all species were present. Age data in table 2 for the

Southern Appalachians were obtained from the Great Smoky Mountains National Park

(Blozan 1994) and the Chattooga River Gorge watershed (see footnote 7) that

encompasses the national forests in Georgia, North Carolina, and South Carolina.

42
White, David L.; Lloyd, F. Thomas. 1998. An old-growth definition for dry and dry-mesic oakpine forests. Gen. Tech. Rep. SRS-23. Asheville, NC: U.S. Department of Agriculture, Forest

Service, Southern Research Station. 42 p.

Dry and dry-mesic oak-pine forests are widely distributed from New Jersey to Texas, but

representative old-growth stands are rare. Historical accounts of composition, along with

information from existing old-growth stands, were used to characterize this type. Shortleaf pine

and white oak were the most widely distributed trees across all old-growth stands. Shortleaf was

the primary pine species of the oak-pine type in the Piedmont, Gulf Coastal Plain, and Interior

Highlands, while pitch pine was predominant in the mountainous regions. Scarlet and chestnut

oaks were more prevalent in the Appalachians. Maximum age for pitch and shortleaf pines

exceeded 200 and 300 years, respectively. Forests of this type do not adhere to the steady state or

equilibrium concept of forest dynamics; rather old-growth oak-pine may be ephemeral on a given

site. Disturbance by fire or other agents distributed through space and time is required for its

maintenance on the broader landscape. Characteristics of the living and dead components of old-

- growth oak-pine are presented.

Keywords: Disturbance, dry-mesic, fire, oak-pine, old growth, presettlement.
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