___________

The Author:

___________

William R. Harms, Research Forester, Southern Research Station,

U.S. Department of Agriculture, Forest Service, Charleston, SC 29414.

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-growth resource. 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 between 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 1996

Southern Research Station

P.O. Box 2680

Asheville, NC 28802
An Old-Growth Definition for Wet Pine Forests,

Woodlands, and Savannas

William R. Harms

Introduction

Wet pine forests, woodlands, and savannas of the

Southeastern United States are subtypes within the pine

flatwoods forest region of the Atlantic and Gulf Coastal

Plains. This review briefly describes the edaphic and

vegetation characteristics of the dominant plant communities

in this type group and provides a working definition that may

be used to identify and evaluate stands for inclusion in an

old-growth management program. The old-growth

definitions offered below are based on available published

information on the type group; no field investigations were

made.

The distinction among the vegetation structure of forests,

woodlands, and savannas is determined by the relative

density of the dominant stand and the composition of the

shrub and herb layers. Forests have closed stands of trees

with touching crowns, well-developed understories of trees

and shrubs (if not frequently burned), and little if any grass

or herb ground cover. Woodlands have fewer trees per unit

area than forests, crowns of the dominant stand do not

generally touch, and the understory is dominated by woody

shrubs, giving the woodland a two-layered structure.

Savannas tend to have widely scattered trees and a

predominantly grass-herb understory of high-species

diversity (Christensen 1988, Harcombe et al. 1993, Peet and

Allard 1993).

Forest Type Group Narrative

Wet pine forests, woodlands, and savannas (type no. 27) are

classed as palustrine forests with frequent, low-intensity,

widespread fires. They are found in the coastal plain

physiographic province from Delaware south to Florida and

west to eastern Texas, primarily in the flatwoods of the outer

marine terraces. Representative sites include boggy

nonriverine flatlands, poorly drained seasonally wet coastal

flatlands, and lowlands adjacent to ponds, streams, and other

wet areas. Soils range widely in texture from clay to sand.

Moisture conditions are influenced by impermeable soil

layers that restrict the downward movement of water. Poor

drainage limits aeration of the soils on many sites and peat

mats commonly develop over mineral surfaces. Acidic

conditions prevail on most sites.

Three principal tree species are characteristic of the wet

pine type group: longleaf pine (Pinus palustris Mill.), slash

pine (P. elliottii Engelm), and pond pine (P. serotina

Michx.). They define specific subtypes and occur either in

pure stands or in mixtures, depending on geographical

location, soil and hydrological conditions, and fire regime.

For ease of reference, the subtypes are named according to

the forest cover type classification system published by the

Society of American Foresters (SAF) (Eyre 1980). The

types are wet-site variants of longleaf pine (SAF 70),

longleaf pine-slash pine (SAF 83), slash pine (SAF 84), and

pond pine (SAF 98).

Species composition differs widely among subtypes and is

shaped by fire frequency and intensity, soil characteristics,

season and duration of flooding or soil waterlogging, and

latitude. The subtypes are segregated on the landscape

primarily on the basis of the soil moisture-hydroperiod

gradient. On mineral soils where flooding is limited,

longleaf pine is the predominate overstory tree. The stands

tend to be pure with few if any trees in the midstory. Fires

on longleaf sites occur at 2- to 5-year intervals. On wetter

sites where fire is less frequent, both longleaf and slash pine

dominate the overstory. On sites with growing-season

hydroperiods, slash pine dominates. Fire usually restricts

hardwood associates, such as sweetgum (Liquidambar

styraciflua L.), red maple (Acer rubrum L.), blackgum

(Nyssa sylvatica Marsh.), and cabbage palmetto [Sabal

palmetto (Walt.) Lood. ex J.A. & J.H. Shult], to understory

positions on sites where they occur. On organic soils

subject to prolonged flooding, longleaf and slash pine are

replaced by pond pine communities. Associates of pond

pine include swamp tupelo [N. sylvatica var. biflora (Walt.)

Sarg.], water oak (Quercus nigra L.), baldcypress

[Taxodium distichum (L.) Rich.], pondcypress (T.

ascendens Brongn.), sweetbay (Magnolia virginiana L.),

live oak (Q. virginiana Mill.), loblolly-bay [Gordonia
2

lasianthus (L.) Ellis], and redbay [Persea borbonia (L.) terraces in southwest Louisiana and southeast Texas

Spreng.]. (Harcombe et al. 1993).

Fire

All the communities in this type group are fire dominated

and depend on fire for their continued existence (Christensen

1981). Although the frequency and intensity of fire vary

among the different subtypes, it is a natural phenomenon in

each one. Without continued disturbance from fire, it is

likely that these vegetation types would dominate a particular

site for a single generation, then gradually lose control to

invading species and eventually disappear from the

community (Wade 1983). The pre-settlement natural fire

regimes for these ecosystems are not known, but studies of

species life histories and patterns of fuel accumulation

indicate that light-to-moderate intensity surface fires

occurred every 2 to 8 years in the grass- and herb-dominated

savannas (Wahlenberg 1946, Christensen 1981). In the

wetter, shrub-dominated slash pine and pond pine

woodlands, fire frequency was probably 10 to 30 years, less

than in the savannas, because of the more moist conditions.

Frequency in these types depends on occurrence of drought

conditions sufficient to increase the flammability of the

understory to where it will burn readily. Such fires are

intense and usually burn all the aboveground vegetation,

especially in pond pine woodlands (Christensen 1981, 1993).

Fire is necessary for the establishment and maintenance of

these ecosystems. In the longleaf types, grasses, such as

wiregrass, are essential fuel components to ensure that fire

will occur frequently (Noss 1989). The range of ecological

processes and conditions that recurring fire initiates,

terminates, and continues cannot be duplicated by any other

disturbance (Volga 1979).

Forest Subtype Narratives

Longleaf Pine Type

Range—The natural range of longleaf pine covers most of

the coastal plain from southeastern Virginia south to central

Florida and west to Texas. It also occurs in areas of the

piedmont and interior uplands of Georgia and Alabama

(Wahlenberg 1946). Wet longleaf pine sites occur primarily

in the flatwoods section of the Atlantic and east Gulf Coastal

Plains and on seepage areas in the fall line section from

central North Carolina south to Alabama (Peet and Allard

1993). In the west Gulf region, wet pine lands occur on

poorly drained interstream areas of the outer coastal plain

Sites—Wet pine sites are seasonally wet, often saturated

during the winter and droughty during the growing season.

Most soils are sandy with low organic matter content and

include predominantly Aquods (Spodosols) and Aquults

(Ultisols). They are characterized by high and fluctuating

water tables, often with a fine textured spodic (organic) or

argillic (clay) horizon that restricts drainage (Boyer 1990).

Vegetation—High stand densities are not typical of

longleaf pine in the natural state (Landers et al. 1990);

virgin stands are described as mosaics of widely scattered

trees intermingled with patches of seedlings, saplings, and

pole-sized trees, and an understory of grasses and herbs

(Schwartz 1907, Wahlenberg 1946). Longleaf pine

communities, therefore, are typically woodlands or

savannas. Longleaf is pure or is the dominant species on

poorly to moderately drained soils. On wetter soils from

southern South Carolina south and west along the Gulf,

slash pine is a common associate or may dominate. In

Florida and southeast Georgia, slash pine and/or pond pine

replace longleaf pine on wet sites. Pond pine assumes

dominance in Virginia and the Carolinas on the wettest sites

and on organic soils. 

In woodland stands, the understory is absent or may contain

invading hardwoods such as blackgum, sweetgum, and

water oak. A low shrub layer of varying density is usually

present. Common species include gallberry [Ilex glabra (L.)

Gray], redbay, sweetbay, and swamp cyrilla (Cyrilla

racemiflora L.). On better drained sites, saw-palmetto

[Serenoa repens (Bartr.) Small] may dominate the shrub

layer. The herb layer is dominated by wiregrass (Aristida

stricta L.). On wet savannas, the dominant pines occur in

stands of widely scattered trees. Understory trees and

shrubs are generally absent. Scattered gallberry, blueberry

(Vaccinium spp.), waxmyrtle (Myrica cerifera L.), or other

shrubs may be present. The herb layer is generally dense

and is very diverse, with grasses, sedges, composites,

orchids, and lilies particularly prominent. In the eastern part

of its range and south to central Alabama and northwest

Florida, wiregrass is the dominant species. In the western

part of the range, bluestem (Andropogon L. spp.) and panic

(Panicum spp.) grasses are dominant (Boyer 1980, Frost et

al. 1986, Harcombe et al. 1993, Peet and Allard 1993).

Community dynamics—Longleaf pine is the longest lived

species of this type group; Platt et al. (1988) reported trees

older than 400 years. The longleaf pine types are fire

maintained successional stages in the natural ecological
3

progression to the southern mixed hardwood climax forest associated species may include swamp tupelo, sweetbay,

(Quarterman and Keever 1962). The existence of pure pine pondcypress, loblolly-bay, live oak, pond pine, southern

stands in the virgin forest is attributed to frequent, naturally redcedar [Juniperus silicicola (Small) Bailey], cabbage

occurring, low-to-moderate intensity surface fires. Such fires palmetto, red maple, and, less commonly, sweetgum, water

create a favorable environment for seedling establishment oak, and swamp laurel oak (Q. laurifolia Michx.).

and growth by keeping competing shrubs and herbs and Seasonally flooded bays dominated by slash pine will have

invading hardwoods in check and controlling the brown-spot sweetbay or loblolly-bay and sometimes swampbay [P.

needle blight [Scirrhia acicola (Dern.) Siggers], which is a borbonia var. pubescens (Pursh) Little], pondcypress, and

major cause of poor seedling growth and mortality (Boyer swamp tupelo in the canopy. The understory will often

1990). Frequent fire is essential for maintaining species include swamp cyrilla, and buckwheat-tree [Cliftonia

diversity and community structure in the longleaf pine types monophylla (Lam.) Britton ex Sarg.]. The herb layer is

(Landers et al. 1995). Without frequent fire, the fire-sensitive absent except in bays where sphagnum species are present.

shrub and hardwood components already present in the On boggy flatwoods sites, slash pine is commonly mixed

stand, plus invaders from seed sources outside the stand, with longleaf pine and a shrub layer of waxmyrtle,

increase and eventually suppress the pines and dominate the gallberry, buckwheat-tree, dahoon (I. cassine L.), and

site (Wahlenberg 1946, Quarterman and Keever 1962). yaupon (I. vomitoria Ait.) is usually present. Species of

Slash Pine Type

Range—Slash pine has the most restricted natural range of lived species; mature trees more than 100 to 150 years old

the major southern pines. It is native only on a narrow strip have seldom been reported (Mohr 1897, Pomeroy and

of flatwoods along the southeast coastal plain from southern Cooper 1956). The slash pine type is fire maintained but,

South Carolina into central Florida and westward along the because seedlings are less tolerant of fire than longleaf pine,

Gulf Coastal Plain to eastern Louisiana. Its range has been it is generally restricted to wetter, less frequently burned

extended by planting west through central Louisiana into sites (Abrahamson and Hartnett 1990). It is a transition

eastern Texas, north in Alabama, Mississippi, and southern species lying on a soil moisture gradient between seasonally

Arkansas, and in parts of North and South Carolina (Hodges wet longleaf pine sites and seasonally flooded ponds and

1980, Lohrey and Kossuth 1990). bay swamps. With the elimination of fire, succession of

Sites—Slash pine occupies wetter sites than longleaf. The

original distribution of the species within its natural range

was largely determined by its susceptibility to fire injury

during its seedling stage, and its tolerance of wet soil Longleaf Pine—Slash Pine Type

conditions (Lohrey and Kossuth 1990, Stout and Marion

1993). It grows in irregular stands often mixed with longleaf Range—The geographic range of this cover type is

or loblolly pine (P. taeda L.), pondcypress, swamp tupelo, determined by the natural range of slash pine. It extends in

and other wetland species. Historically, it was not noted for the coastal plain flatwoods from southern South Carolina to

forming large forests, and extensive natural stands occurred southeast Louisiana. 

only in central Florida (Mohr 1897, Schultz 1983). The sites

where slash pine grows naturally include wet savannas and

pitcher plant flats, poorly drained flatwoods, stream edges

and pond margins, and seasonally flooded areas such as

bays, ponds, and swamps where fires are rare (Mohr 1897,

Hodges 1980, Schultz 1983). The soils range from

Ochraquults and Albaquults (Ultisols) to Humaquents

(Entisols), depending on the surface texture, color, and depth

to a clay layer. They vary from loamy sands to sandy loams

in surface texture. Because they are wet, they often

accumulate organic matter (Pritchett and Comerford 1983).

Vegetation—The community has an open-to-closed tree

canopy comprised predominantly of slash pine. On wet sites,

pitcher plants occur in the herb layer (Hodges 1980). 

Community dynamics—Slash pine is a relatively short

wet-site slash pine tends toward the bay community type

(Monk 1968) or swamp hardwoods (Abrahamson and

Hartnett 1990).

Sites—Fire history and moisture regime of the site are

responsible for the occurrence of this type. It occurs on

flatwoods and savanna sites where longleaf pine is adjacent

to or in the vicinity of a slash pine seed source. In the

prolonged absence of fire, slash pine will seed into

neighboring longleaf stands to become established in the

understory and, over time, grow into the overstory. Because

slash pine seedlings are susceptible to fire damage, the site

must be fire-free for 5 to 10 years to allow seedlings to

reach fire-resistant size of approximately 9.8 feet (3 meters)

tall (Grelen 1980).
4

Vegetation—Slash and longleaf pines occur in the overstory

in varying proportions. The most common shrub associates

are gallberry and saw-palmetto. On wet sites, sweetbay,

blackgum, buckwheat-tree, titi, greenbriers (Smilax spp.),

fetter-bush [Lyonia lucida (Lam.) K. Koch], sweet

pepperbush (Clethra alnifolia L.), St. Andrew’s Cross

(Ascyrum hypericoides L.), and blueberries are common.

Wiregrass is a major component of the herbaceous layer if

fire is frequent enough to control the abundance of the shrub Community dynamics—Pond pine communities are wet

layer (Grelen 1980). and nutrient-poor and are susceptible to fires during

Community dynamics—With or without fire, this

community is temporary. Fire destroys young slash pine; but

where exclusion of fire allows slash pine to become

established with longleaf pine, hardwoods and shrubs will

invade and dominate the understory and eventually the

overstory, eliminating subsequent regeneration of both pines

(Grelen 1980). With frequent fire, the type will revert to

longleaf pine. 

Pond Pine Type

Range—Pond pine occurs in the coastal plain from southern

New Jersey south to central Florida and west into

southeastern Alabama (Bramlett 1990).

Sites—Pond pine occupies the wettest sites within this type

group. It is found in swamps, pond borders, and pocosins,

characterized by undrained peat soils and wet, sandy flats. It

grows most extensively in the broad, poorly drained

interstream areas of peaty soils in the lower coastal plain of

North Carolina. Farther south, it is found on the wet pine

flatwoods and savannas of the lower coastal marine terraces

and in bays and ponds throughout the coastal plain. The

species can make excellent growth on better drained mineral

soils, but the faster earlier growth of slash and loblolly pine

usually restricts it to a subordinate position on such sites

(Johnson 1980, Bramlett 1990). The soils range from

Ochraquults and Albaquults (Ultisols) to Humaquents

(Entisols), depending on the surface texture, color, and depth

to a clay layer. Organic soils include oligotrophic (nutrientpoor, rain-fed) mineral soils with shallow organic surface

layers, represented by Haplaquods (Ultisols), and shallow-todeep Humaquepts and Medisaprists (Histosols) (Schafale

and Weakley 1990).

Vegetation—The dominant tree canopy of pond pine stands

is open woodland to nearly closed forests. Associates include

sweetbay, swampbay, red maple, loblolly pine, and Atlantic

white-cedar [Chamaecyparis thyoides (L.) B.S P.] The shrub

layer includes swamp cyrilla, fetter-bush, maleberry [L.

ligustrina (L.) DC], gallberry, blue huckleberry [Gaylussacia

frondosa (L.) T.&G.], saw-palmetto, sweet pepperbush,

laurel-leaved greenbrier (S. laurifolia L.), and swampbay.

Switch-cane (Arundinaria spp.) is often present and may

dominate in Atlantic Coastal Plain sites. Saw-palmetto is

the dominant shrub in Georgia and Florida. There is

generally no herb layer (Edmisten 1965, Schafale and

Weakley 1990).

droughts that allow the organic soils to dry. The large

amount of fuels produced by the understory make fires

extremely intense. Pond pine has serotinous cones that

require hot fires to release seed and produces epicormic

sprouts when injured. Regeneration of new stands is

therefore tied directly to fire (Woodwell 1958). All the

dominant species sprout readily and the shrub layer regrows

to its former height in just a few years. Stands persist

indefinitely in the absence of fire but are readily regenerated

after burning by sprouting, as well as from seeds (Schafale

and Weakley 1990).

Old-Growth Definitions

Elimination of fire regimes of natural and Indian origin,

indiscriminate logging, and conversion of forests to farms

produced sweeping changes in the vegetation landscape that

was present in the Southeast at the time of European

settlement. As a result, few stands representative of the

virgin or old-growth conditions of wet pine forests still

exist. New old-growth forests will have to develop from

existing stands with the help of an active management

program and a set of old-growth definitions to guide it.

Published data from examples of virgin or old-growth

forests of this type group from which old-growth definitions

can be constructed are limited. There are, however,

descriptions of upland longleaf stands (Chapman 1907,

Schwatrz 1907, Wahlenberg 1946, Platt et al. 1988) and a

slash pine stand (Hebb and Clewell 1976) that provide

limited guidance. No descriptions of old-growth pond pine

have been found.

Provisional Definitions

These definitions are provisional in that they are based on

limited information. They should be modified as new

information from research and field study becomes

available. There are five basic stand-selection criteria for

the wet pine forests, woodlands, and savannas: (1) The

environment must have seasonally wet soils. (2) It must
5

include the fire regime appropriate to one of the subtypes

described. (3) There must be a stand of trees of the required

species composition and minimum age. The minimum age of

most stands in all subtypes in this group will be 80 to 100

years, assuming that most of the oldest stands date from the

late 1800’s to the early 1900’s, when the last of the oldgrowth southern pines were harvested (table 1). (4) There

should be no evidence of recent soil disturbance or tree

harvest. (5) Stand structures must be appropriate for forest,

woodland, or savanna. Given these criteria, other

characteristics and attributes of old-growth forests, such as

snags, large and small woody debris in various stages of

decay, herbaceous and woody undergrowth typical of the

type, and regeneration of the dominant canopy species must

be either present or the conditions necessary for their

development evident.

Stand attributes used to identify old-growth wet pine forests

are presented in table 1. Standards are defined for each of the

forest subtypes except for longleaf pine-slash pine.

Standards for this type are assumed to be the same as for the

individual longleaf and slash pine types. Decadence in old,

dominant trees, such as dead, broken, or deformed tops, and

stem or root rot, also indicates old growth and should be

evaluated. The minimum area for a stand to be considered

viable for old growth is not defined. Acceptable size must be

determined in the context of the landscape in which the stand

occurs; some factors that should be considered when

determining stand size include extent of edge effects from

neighboring stands and ecosystems, and vulnerability to

catastrophe.

Table 1—Provisional standards for old-growth wet pine

forests, woodlands, and savannas in the southeastern

coastal plaina

Standard attributesb

_____________________________________________________________

Minimum Average Minimum

Species no. trees/ha stand d.b.h. age

cm yr

Live canopy trees

 (forest structure):

Longleaf pine 150 50 150 - 200

Slash pine 150 53 80 - 100

Pond pine 200 23 60 - 100

Stand structure :

c

Forest Hardwood understory present

Woodland Shrub understory dominant

Savanna Grass-herb understory dominant

 Forest = most crowns touching or nearly touching; woodland = trees a

widely scattered, shrub layer prominent; savanna = few scattered trees,

herb layer prominent.

 Standards are for forest conditions and are extracted from various b

sources: longleaf pine (Chapman 1907, Wahlenberg 1946); slash pine

(Hebb and Clewell 1976); pond pine—extrapolated from natural stand

yield tables of Schumacher and Coile (1960) for age 80 and site index 70.

Standard attributes for canopy trees in woodlands and savannas are not

known—they can be set using the forest standards as a base.

Stand structure will depend on the existing fire regime. Frequent fires c 

favor a grass-herb understory, while less frequent fires favor a hardwood

understory.
6

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7

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Woodwell, G.M. 1958. Factors controlling growth of pond pine seedlings

in organic soils of the Carolinas. Ecol. Monogr. 28:219-236.
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forests, woodlands, and savannas. Gen. Tech. Rep. SRS-2. Asheville,

NC: U.S. Department of Agriculture, Forest Service, Southern

Research Station. 7 p.

The ecological, site, and vegetation characteristics of pine wetland

forests of the flatwoods region of the Southeastern United States are

described. Provisional working definitions of old-growth characteristics

are provided for longleaf pine, slash pine, and pond pine forests. These

definitions can be used to identify and evaluate stands for retention in

old-growth management programs.

Keywords: Forested wetlands, longleaf pine, old growth, old-growth

management, pine flatwoods, pond pine, slash pine.

.
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