The Authors:

J. Larry Landers, Deceased, Joseph W. Jones Ecological Research Cerner

at Ichauway, Newton, GA 3 1770; and William D. Boyer, Retired, Southem

Research Station, U.S. Department of Agricultnre, Forest Service,

Aubum University, Aubum, AL 36849.

Preface

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

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

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

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

contribution to biodiversity were apparent. It was also evident that definitions were needed to adequately assess sud 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 enhancmg 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 tmiformity 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, al1 Forest Service Stations and Regions began developing old-growth defmitions

for specific forest types. Because the Southem and Eastem Regions share many forest communities (together they encompass the

entire Eastem United States), their efforts were combined, and a cooperative agreement was established with The Nature

Conservancy for technical support. Tbe resulting project represents the tirst large-scale effort to define old growth for al1 forests

in the Eastem United States. This project helped briug the old-growth issue to public attention in the East.

Deíinitions 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 deftitions will be drafted.

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

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

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

organizational and geographic boundaries.

August 1999

Southern Research Station

P.O. Box 2680

Asheville, NC 28802
An Old-Growth Definition for Upland

Longleaf and South Florida Slash Pine

Forests, Woodlands, and Savannas

J. Larry Landers and Wi.Iliam D. Boyer

Introduction

Longleaf pine (Pinuspalustris Mill.) and South Florida slash

pine (l? elliottii var densa Little & Dorman) ecosystems are

distinguished by open, multiaged forests, woodlands, or

savannas, sometimes referred to as the “pine barrens.”

Longleaf pine once occupied the largest area dominated by a

single-tree species (Chapman 1932, Frost 1993) in North

America. Although longleaf is a very intolerant pioneer

species, it was ecologically persistent rather than

successional or transitional (Platt et al. 1988), and

maintained itself in place for thousands of years. The

southeastem pine belt is noted for the high Cequency of

thunderstorms that generated the most chronic fire regime of

any forest region in North America. The traits of these two

pine species and associated native grasses played a key role

in the facilitation of fue, an enviromnental agent essential to

the competitive success of these trees and the maintenance

of their remarkably diverse, fue-dependent ecosystems. The

old-growth setting for both longleaf and South Florida slash

pine, which resembles longleaf within its limited range, is

best characterized as a rich ensemble of grasses and other

herbaceous vegetation topped only by pines of variable ages

and densities.

This document describes probable old-growth conditions for

upland longleaf and South Florida slash pine forests,

woodlands, and savannas based on currently available

information. Forests, woodlands, and savannas as defined

here follow a standard terminology (Ford-Robertson and

Winters 1983). A forest is an ecosystem characterized by a

more or less dense and extensive tree cover; a woodland is a

plant community in which trees are present but form an open

canopy with intervening areas occupied by lower vegetation,

commonly grasses; and a savanna (tree savanna) is generally

a grassland with a scattering of trees and shrubs. The

definitions are based on general appearance of the land

rather than on numbers of trees or tree basal area per unit of

area.

Description of Upland Longleaf

and South Florida Slash Pine

Forests, Woodlands, and Savannas

Range

The combined mnges of longleaf and South Florida slash

pine (fig. 1) once extended across most of the southeastem

Coastal Plain and southem reaches of the adjacent Piedmont,

Blue Ridge, and Valley and Ridge physiographic provinces

(Fenneman 1938). Longleaf pine-dominated communities

ranged fiom southeastem Virginia through most of Florida

and into eastem Texas (Critchfield and Little 1966, Frost

1993). South Florida slash pine occurs exclusively in central

and southem Florida where it is considered the ecological

equivalent of longleaf pine (Mohr 1897).

soils

Three soil orders are of major importance in this pine region

(U.S. Department of Agriculture 1969). Ultisols are the

principal order over much of the Deep South outside of

peninsular Florida. Ultisols most commonly associated with

longleaf are the Typic and Plinthic Paleudults. Deep, sandy

Entisols, which are primarily Quartzipsamments, range fiom

about 10 feet [3 meters (m)] above sea leve1 in Florida up to

600 feet (183 m) in Georgia and the Carolinas; they form the

sandhills of the Carolinas, Georgia, and northwestem Florida

and the sand ridges in the central highlands of Florida.

Surface soils are typically acidic and low in organic matter,

nitrogen, calcium, and phosphorus (Wahlenberg 1946). On

the Coastal Plain, these pines grow in sandy (and some

loamy or clayey) surface soils on well-drained rolling or

sloping sites, across dry to xeric hills or ridges, and on rises

within moist flatwoods. In many areas they gradually incline

to boggy flatwoods or wetland depressions. Lower soil

horizons vary from almost pure sand to clays. Within

mountain provinces, longleaf occupies thin, stony soils on

dry ridges and slopes with South-to-southwest exposures.

The South Florida slash pine is commonly found on thin

soils over lime rocklands.
Figure 14eogmphic mnges of longleaf pine (light gtay) and South Florida slash pine (dark gmy) (Critchfield and Little 1966).

2
Trees Understory Vegetation

Dominant longleaf or South Florida slash pines typically

occur in ahnost pure stands, often with patchy to open

canopies. Clusters of deciduous scrub oaks-turkey oak

(@emus luevis Walt.) and/or bluejack oak (Q. incana Bart.)

on xetic or subxeric sands, post oak (Q. margaretta Ashe)

on loamy sands, and blackjack oak (Q. marilandica

Muench.) on clayey soils-sometimes occur. Some evergreen

oaks, e.g., sand live oak (Q. geminatu Small), may be

significant natural associates of longleaf pine in coastal

scrub woods. They are more tolerant of fire than other

hardwoods. Some can grow into the midstory despite

biennial, dormant-season fires (Boyer 1993).

Evergreen oak scrub or sand pine [P clausa (Chapm. ex

Engehn.) Vasey ex Sarg.]-evergreen oak scrub communities

in some partially fue-protected areas may invade open pine

communities where the fue regimes have been disrupted.

Similarly, an abundance of mesic hardwoods growing with

longleaf suggests an extended period of fire exclusion,

overcuttmg, or soil disruption (Wahlenberg 1946). A mixture

of both longleaf and slash pine, a recognized forest type,

may also be a transition commuuity reflecting a long-term

change in fire intervals. Some early observations have

suggested that mixtures of longleaf, loblolly (I? taedu L.),

and shortleaf (P. echinatu Mill.) occurred with various mesic

hardwoods on some hilly sites (Frost 1993); but few natural

mixtures have been documented. Some associations were

probably similar to those of southwestem Georgia where

longleaf pine, oak, and hickory (Cava spp.) occurred at a

ratio of 9 1: 1 :OS at the time of European settlement (1733-

1832) (Plummer 1975). Other communities with a large

hardwood component in the overstory and midstory are

probably in transition f?om pine to late-successional, mixedhardwood forest. Longleaf pine cannot succeed itself under

these conditions, and the transition will be complete when

the last of the residual pines die.

South Florida slash pine, also a fue-tolerant species, is ofien

the sole canopy tree in woodlands and savarmas, except in

southcentral Florida where it overlaps with longleaf. On

some high-quality sites, low densities of cabbage pahn

[Sabalpalmetto (Walt.) Lodd. ex Schult. & Schult.], which

also tolerates frequent surface fires (Wade et al. 1980), and

occasionally oaks, e.g., Q. nigru L., Q. virginiana Mill., may

be considered natural associates. In rockland habitats,

numerous broad-leaved subtropical trees, including exotics,

have invaded South Florida slash pine sites. These trees have

made native overstory associations diffrcult to determine.

Both longleaf and South Florida slash pine ecosystems are

named for the dominant and oRen exclusive canopy tree.

Fires that maintained these forests over the millennia also

supported fue-dependent plant communities on the forest

floor. The broad geogmphic range of longleaf pine includes

wide variations in the physical environment based on

physiography, climate, soils, and topographic position. Many

different understory plant community types developed in

response to changing physical environments. The most

distinctive changes within a locale are related to soilmoisture gradients. Twenty-three plant community types

have been identified in longleaf forests, woodlands, and

savannas east of the Mississippi River (Peet and Allard

1993). Further research will surely revea1 many more.

Similarly, a number of understory plant communities have

been recognized in West Gulf longleaf ecosystems

(Harcombe et al. 1993).

Although the ground cover beneath longleaf pine contains

hundreds of herbs and low shtubs, dominant plants normally

are the pyrogenic grasses, important to maintaining these

fue-dependent communities-including the overstory pine.

Wiregrass (Aristida stricta or A. bqrichiana) is of major

importance, particularly east of South-central Alabama and

the westem edge of the Florida Panhandle; bluestem grasses,

e.g., Schizachyrium tenerum, S. scoparius, dominate most

other areas (Wright and Bailey 1982). Abundant wiregrass is

indicative of virgin sites because severe soil disturbance,

such as plowing or intensive mechanical site preparation,

will eliminate this species (Clewell 1989). Certain bluestems

characterize most other locations. There is a similar

relationship between South Florida slash pine and these

same genera of grasses, along with scattered shrub patches

(Snyder 1986). Today, mantles of saw pahnetto (Serenoa

repens) are cornmonly found with South Florida slash pine

(Gunderson and Loope 1982), but this shrub has so

proliferated that its historie range is unknown.

Disturbance Regimes

The natural fire frequency needed to maintain longleaf pine

ecosystems on upland sites is estimated to be 2 to 4 years

(Landers et al. 1990). Fire intervals of 3 to 10 years have

been estimated for longleaf in Coastal Plain sandhills where

edaphic factors reduce the rate of fuel accumulation,

topographic position, or both, limits fue entry (Wharton

1978, Clewell 1986) in the species’ “transition” region and

in other partiahy fire-protected areas (Ware et al. 1993).

Before the natural systems were disrupted, fire-dependent

3
South Florida slash pine communities were similarly

maintained by regularly recurring surface fires at 2- to 7-

year intervals (Harper 1927, Wade et al. 1980, Gtmderson

and Loope 1982, Snyder 1986).

Far-ranging lightning Eres were common until forest

fiagmentation became widespread and forest fuesuppression policies were established (Robbins and Myers

1992). Komarek (1964) found that about 93 percent of

lightning ignitions in northern Florida occurred fiom late

May to early August. Most lightning fires occurred in May

and June, although thunderstorms were most frequent in July

and August. For south Florida, Snyder (1986) detected

similar seasonal wildfire pattems.

Catastrophic events other than lightning fires occur within

normal tire regimes-blowdowns, intense fues following

prolonged droughts, and subsequent beetle attacks-that can

variously impact pine forests (Schwarz 1907, Platt and

Rathbun 1993). Widespread droughts of 5 or more years

occur about every 22 years and altemate with periods of

abundant rainfall (Plummer et al. 1980). Most of this region

will experience tropical storms every one or two decades

(Neumann et al. 1987, Hooper et al. 1990); one or two

hurricanes are likely to make landfall every year. Tomados

and microbursts are also common throughout most of the

South.

Crosswalk with Society of Ameritan Foresters Forest

Cover Qpes (Eyre 1980)

70-longleaf pine

7 1-longleaf pine-scrub oak

83-longleaf pine-slash pine

ll l-South Florida slash pine

Physiographic Provinces (after Fenneman 1938)

l Coastal Plain (al1 sections)

l Piedmont (upland section)

l Blue Ridge (southem section)

l Valley and Ridge (southem section)

Old-Growth Conditions

Today only a few old-growth longleaf and South Florida

slash pine communities remain, and these fragments may not

be large enough or ecologically cohesive enough to be selfsustaining over the long term. Only sketches of old-growth

conditions can be assembled from reported observations, old

photographs, and the few studies of surviving relicts. Al1 of

these information sources were used to characterize oldgrowth attributes over the ranges of forests, woodlands, and

savannas that were once dominated by these two species.

Old-growth characteristics are summarized in table 1 along

with the source cited.

Living Tree Component

F. Harper (1942) and M. Van Doren (1940) described sparse

longleaf woodlands or savannas on the wettest and driest

sites, and woodlands or forests of large, dense pines on

intermediate sites. F. Harper (1942) noted short, large

diameter pines on xeric sands in Florida; and in a savarma he

estimated 20 to 100 yards (18 to 9 1 m) between pines, or

about 0.2 to 3.9 trees per acre [0.5 to 9.5 per hectare (ha)].

Along a fa11 line sand ridge in Georgia, Muir (19 16) also

noted wide spacing of longleaf pines 20 to 30 inches [5 1 to

76 centimeters (cm)] in diameter but only 60 to 70 feet ( 18

to 2 1 m) tall. Spacing in some virgin longleaf woodlands of

southem Georgia was 30 to 50 feet (9.1 to 15.2 m) (Harper

1922-23), i.e., about 6 to 15 large trees per acre (15 to 37

per ha). Some areas in eastem Texas contained 9 to 60 oldgrowth longleaf trees per acre (22 to 148 per ha). They were

100 to 350 years old (Chapman 1909).

Schwarz ( 1907) summarized measurements from six virgin

longleaf stands, one each in Alabama and Mississippi and

four in Louisiana. These were selected as “typical examples

of the character of growth of the main body of virgin

longleaf pine.” Overall, the stands averaged 100 (range 52 to

167) trees per acre [averaged 248 (range 128 to 4 12) trees

per ha] distributed overall in diameter at breast height

(d.b.h.) classes frorn 2 to 34 inches (5 to 86 cm). Stand

density (basal area) averaged a relatively dense 109 square

feet per acre (25 m2 per ha), with a range fiom 68 to 146

square feet per acre ( 16 to 34 m2 per ha) and a mean canopy

cover of 62 percent (range 50 to 80 percent). Virgin stands

were not necessarily old growth. One of Schwarz’s six

stands had three-fourths of al1 stems in the 8- to 12-inch (20-

to 30-cm) d.b.h. classes, and nothing above the 18-inch (46-

cm) class. Another had 82 percent of al1 stems in the lo-inch

(25-cm) and smaller d.b.h. classes, with the largest munber

in the ó-inch (15-cm) class. However, this stand also had

four trees per acre (10 per ha) ranging f?om 20 to 28 inches

(5 1 to 7 1 cm) in d.b.h. The larger trees would seem to have

been older relicts among which a younger stand had grown.

Noting that pine-population structure does not maintain its

uniformity over more than a few hundred acres, often

changing abruptly within 50 acres (20 ha), Schwarz

characterized virgin forest as “. . . fiequent transitions in the

4
Table 1 (English units)-Standardiied table of old-growth attributes for upland longleaf and South Florida slash pine

forests, woodlands, and savannas

.Quantifiable

attribute

Value Number

Range Mean of stands” Referentes

Pine stand density (no./acre)

-treesr4 in. d.b.h. 0.2-100 40 40 Harper 1942

Chapman 1905

Chapman 1909

Doren et al. 1993

Gunderson and Loope 1982

Harper 1922-23

Meldahl et al. 1995

Platt et al. 1988

Schwarz 1907

Pine stand basal area (Wacre)

-trees 9in. d.b.h. 0.2-110 35 38 Harper 1942

Chapman 1905

Chapman 1909

Doren et al. 1993

Gunderson and Loope 1982

Harper 1922-23

Meldahl et al. 1995

Plan et al. 1988

Schwarz 1907

Age of large trees (yr~)~

-1ongleaf pine 100-350 -

-South Florida slash pine

Number of 4-m size classes

lO(r300 -

-1ongleaf trees 22 in. d.b.h. 5-8 7

D.b.h. largest trees (in.)

-1ongleaf pine 19-32 23

-South Florida slash pine 8-12 10

- Chapman 1905

Chapman 1909

Meldahl et al. 1995

Platt et al. 1988

Schwarz 1907

- Doren et al. 1993

7 Chapman 1905

Schwarz 1907

33

1

Chapman 1909

Meldahl et al. 1995

Muir 1916

Platt et al. 1988

Schwarz 1907

Doren et al. 1993
Table 1 (English units)-Standardized table of old-growth attributes for upland longleaf and South Florida slash pine

forests, woodlands, and savannas (continued)

Quantifiable

attribute

Standmg snags (no./acre)

-1ongleaf pine

-South Florida slash pine

Value Number

Range Mean of stand@ Referentes

0.1-7.0 2.7 8 Harper 1942

Engstrom, personal communicationc

Meldahl et al. 1995

Schwarz 1907

3.0-5.8 3.9 1 Doren et al. 1993

Downed logs-longleaf pine

-trees 23 in. (ft3/acre)

-trees 210 m. d.b.h. (no./acre)

- 183

0.4-4.5 2.3

1 Landers, unpublished datad

1 Hermann 1993

No. of canopy layers l-2 - - -

Percent canopy in gaps 20-80 50 8 Harper 1942

Platt, personal communicatione

Schwarz 1907

’ Number of stands may not equal the number of citations.

* Jncludes dominants and codominants in the upper canopy.

’ Personal communication. [Date unknown]. R.T. Engstrom, Wildhfe Biologist, Tall Timbers Research Station, Route 1, Box 678, Tallahassee, FL

323 12.

d Landers, J.L. 1993. Data on longleafpine coarse wood debris in Hauss Park, Flomaton, AL: 183.2 ft3/ac (12.8 mVha) as determined by hne-transect

method. Unpublished data. Gn file with: Joseph W. Jones Ecological Research Center at Ichauway, Newton, GA 31770.

’ Personal communication. 1993. W.J. Platt, Professor, Department of Plant Biology, Louisiana State University, Baton Rouge, LA 70803.
Table 1 (metric units)-Standardized table of old-growth attributes for upland longleaf and South Florida slash pine forests,

woodlands, and savannas

Quantifiable

attribute

Value Number

Range Mean of stands“ Referentes

Pine stand density (no./ha)

-trees 210 cm d.b.h.

Pine stand basal area (m%a)

-trees 210 cm d.b.h.

Age of large trees (yrs)

-1ongleaf pine

-South Florida slash pine

Number of 1 O-cm size classes

-1ongleaf trees 25 cm d.b.h.

D.b.h. largest trees (cm)b

-1ongleaf pine

-South Florida slash pine 20-3 1

0.5-247

0.05-25.2

100-300

5-8

48-82

99 40 Harper 1942

Chapman 1905

Chapman 1909

Doren et al. 1993

Gunderson and Loope 1982

Harper 1922-23

Meldahl et al. 1995

Platt et al. 1988

Schwarz 1907

8.0 38 Harper 1942

Chapman 1905

Chapman 1909

Doren et al. 1993

Gunderson and Loope 1982

Harper 1922-23

Meldahl et al. 1995

Platt et al. 1988

Schwarz 1907

-

- Chapman 1905

Chapman 1909

Meldahl et al. 1995

Platt et al. 1988

Schwarz 1907

- - Doren et al. 1993

7 7 Chapman 1905

Schwarz 1907

59 33

25 1

Chapman 1909

Meldahl et al. 1995

Muir 1916

Platt et al. 1988

Schwarz 1907

Doren et al. 1993

7
Table 1 (metric units)--Standardized table of old-growth attributes for upland longleaf and South Florida slash pine

forests, woodlands, and savannas (continued)

Quantifiable

attribute

Standing snags (no./ha)

-1ongleaf pine

-South Florida slash pine

Downed logs-longleaf pine

-trees 28 cm (m3/‘ha)

-trees 225 cm d.b.h. (no./ha)

No. of canopy layers

Percent canopy in gaps

Value Number

hge Mean of stands”

0.2-17.3 2.7 8

7.4-14.3 3.9 1

- 12.8 1

l-ll 5.7 1

l-2 -

-

Referentes

Harper 1942

Engstrom, personal communicationc

Meldahl et al. 1995

Schwal-z 1907

Doren et al. 1993

Landers, unpublished datad

Hermann 1993

-

20-80 50 8 Harper 1942

Platt, personal communication’

Schwarz 1907

0 Number of stands may not equal the number of citations.

* Includes dominan& and codominants in the upper canopy.

c Personal communication. [Date unknown]. R.T. Engstrom, Wildlife Biologist, Tall Timbas Research Statio~ Route 1, B& 678, Tallahassee, FL

32312.

d Ianders, J.L. 1993. Data on longleaf pine toarse wood debris in Hauss Pa&, Flomaton, AL: 183.2 A-‘/ac (12.8 m3/ha) as determined by lin&ransect

method. Unpublished data. On fíle withz Joseph W. Jones Ecological Research Center at Ichauway, Newton, GA 3 1770.

e Personal communication. 1993. W.J. Platt, Professor, Department of Plant Biology, Louisiana State University, Baton Rouge, LA 70803.

8
ages and in the density of growth, so that the effect is often

that of a succession of large and small groves, either

gradually merging into one another or changing abruptly

f?om young and dense to mature and opener growth, orto a

mixture of variable sizes and ages.”

In an analysis of the Wade Tract, an old-growth longleaf

pine preserve in southwestern Georgia, Platt et al. (1988)

showed a rotated sigmoid diameter distribution representing

many loosely aggregated age groups. Mature trees tended

toward random dispersion; the authors estimated about ll

longleaftrees per acre (27 per ha) 100 to 250 years old and

59 younger trees 2 1 inch (2.5 cm) in d.b.h. per acre (146

per ha). The tract included many small gaps, some with

patches of pine regeneration of various ages and others

remaining treeless for years. They concluded that natural

longleaf stands are mrely stable, yet they persist as long as

frequent fires can maintain openings until colonization.

An uncut, old-growth stand in south Alabama has been

degraded due to absence of fne for over 45 years. As a

res&, no longleaf has been recruited, although other pines

and hardwoods have invaded the stand (Meldahl et al.

1995). Longleaf pines were dominant with 128 stems per

acre (3 16 per ha) and a basal area of 82 square feet per acre

(19 m2 per ha). Other trees constituted 1,O 18 stems per acre

(2,5 15 stems per ha) and had a basal area of 44 square feet

per acre (10 mz per ha). Al1 trees > 15 inches (38 cm) in

d.b.h. were longleaf pine. The older trees measured ranged

from 140 to 210 years.

For an old-growth South Florida slash pine tract that had not

been protected fiom wildfires (mostly from April to June),

Doren et al. (1993) documented a similar population

structure. Early narratives about old-growth South Florida

slash pine suggest that there were open, uneven-aged

savannas (Doren et al. 1993).

The approximate age at which old-growth features begin to

appear is about one-half the maximum age of the

predominant tree species (Maine Critical Areas Program

1983). Applying that guideline to average lifespan estimates

for longleaf and South Florida slash pines (Landers 199 1)

suggests that those conditions would appear at about ll 5

and 1 OO years, respectively. Longleaf can be considered

fully mature and perhaps entering old-growth status when

height growth ends, an age that will vary among individuals.

However by 100 to 110 years, height growth has stopped in

about one-half of observed longleaf pines.

Longleaf pine may live 500 years or longer, although

lightning strikes, intense fire, and windthrow generally

shorten its lifespan. Annual longleafattrition of about one

canopy tree per 5 acres (2 ha) in mature, well-stocked stands

(Bayer 1979) suggests that, in the absence of catastrophic

events, the last tree in a cohort would die by age 500.

In medium- to well-stocked, even-aged longleaf pine stands,

a steady state (growth and mortality in balance), which is

another indicator of old-growth conditions, can be expected

to begin at about age 120. Mortality will include normal

attrition among canopy trees and self-thinning, which

gradually eliminates suppressed trees. Depending on initial

stocking density and the absence of any catastrophic event,

the steady-state condition may last up to 200 years before

mortality increasingly exceeds growth and the stand begins

to break up. Formerly suppressed trees or established

regeneration may fil1 gaps created by the death of old

residuals.

At the estimated minimum old-growth age of ll 5 years,

longleaf should begin to display some of the features typical

of old-growth conditions-large boles, contorted and flattopped crowns with a few massive primary branches, decay

within some live trees, near cessation of height growth, resin

core formation, and resinous large woody debris (Chapman

1905, Chapman 1909, Platt et al. 1988). Similar features

have been observed in old-growth South Florida slash pines

(Doren et al. 1993), and the age of some old-growth stands

has been estimated at nearly 200 years (Mohr 1897)

although some stands may reach up to 300 years (Doren et

al. 1993).

A number of pines that are well over a century old would

favor wildlife species whose diet includes pine seeds or who

use large boles for nesting or foraging. A scientific summit

on requirements of the endangered red-cockaded

woodpecker set a target age of 100 to 250+ years for

longleafto provide suitable nesting habitat (Southeast

Negotiation Network 1990). Considering the important oldgrowth structural features and the sensitive ecosystem

elements associated with its structure, a minimum age of 115

years seems an appropriate target for the dominant pine

component of an old-growth longleaf stand.

Sizes of pines in the older age classes are variable because

diameter is strongly influenced by stand density and height

by site quality. Age, as well as the present and former

competitive position in the stand, also affects both diameter

and height of individual trees. At age 115 years, Schwarz’

(1907) longleaf diameter growth curves derived for each tree

in a %-acre (0.2~ha) old-growth stand show a range fiom

about 8 to 25 inches (20 to 64 cm) (scaled from curves) and

average about 16.8 inches (42.7 cm) in d.b.h. When these

9
measurements were made, the largest tree was 2 10 years old,

the smallest was 120 years old. The d.b.h. of the oldest

cohort alone ranged t?om 24 to 32 inches (61 to 8 1 cm). At

age 115, the d.b.h. of the oldest cohort ranged from 17 to 25

inches (43 to 64 cm). Continuing competition from older

trees in the stand slowed long-term diameter growth of the

younger trees. Pine timber 2.20 inches (5 1 cm) in d.b.h.

appears to be necessary to sustain the red-cockaded

woodpecker in the long term (Hamel et al. 1982). Most trees

reaching old-growth status should be near this size or larger.

Dead Tree Component

Snag density, of course, varies with overstory cover, as well

as with disturbarme. Mature, domina& trees are most often

killed by lightning (Komarek 1968, Platt et al. 1988) and

occasionally by intense fire in the growing season, e.g., late

summer (Chapman 1957, Boyer 1990). In turn, canopy trees

often die in chnnps. Schwarz (1907) recorded a mean of six

dead or dying trees per acre (range 0 to 12) [mean of 15

dead or dying trees per ha (range 0 to 30)], representing

mortality over an undetermined number of years. The snags

were mostly small poles but included an average of at least

0.5 large pines per acre (1.2 per ha). Despite a strong taproot

and dense wood, hunicane-forte winds can destroy pine

canopy trees in patches to huge tracts (Schwarz 1907,

Wahlenberg 1946, Bengtson et al. 1993, Platt and Rathbun

1993).

The size of snags associated with old growth is difficult to

surmise because of the great variance in tree size within old

age classes and the variable effects of disturbance. However,

lightning tends to select the largest trees. The minimum

requirements of characteristic wildlife species are usable

indicators of snag size. Of 11 snag-dependent birds in an

old-growth longleaf census by R.T. Engstrom,’ the smallest

species require trees 2 6 inches (15 cm) in d.b.h., and one of

the largest (pileated woodpecker) requires snags 2 20 inches

(51cm) (Hamel et al. 1982).

Snags can stand for many years but are prone to lightning

strikes and susceptible to fnes. Accumulated bark and fallen

branches at the snag base serve as important fuel sources

(Platt et al. 1988). Most early writers described easy travel

through pinewoods, but Cabeza de Vaca ( 1542) noted severe

obstructions due to many downed, lightning-killed trees in

some areas of Florida.

’ Personal communication. [Date unknown]. R.T. Engstroxn, Wildlife

Biologist, Tal1 Timbers Research Station, Route 1, Box 678, Tallahassee,

FL 323 12.

10

Light accumulation of woody debris is normal in old-growth

stands in the South because resin-soaked logs bum readily

and wood decays rapidly. The soil is likely to include fine

charcoal and decaying organic matter from surface deposits

and stump or root charmels. Research is needed to determine

the influente of wood debris on soil characteristics, storage

or flow of nutrients and energy, and facilitation of

decomposers and heterotrophs.

Other

Interactions between fne and downed logs influente plant

community dynamics. During a tire, areas immediately

downwind of logs may become fire shadows where newly

established pine seedhngs may survive (Schwatz 1907).

Burning piles of surface debris often kills all vegetation,

leading to colonization-by including new plants and pine

regeneration. Pine often responds with accelerated grow-th

(the ashbed effect), which also may trigger local changes in

soil chemistry and biota. These processes, coupled with the

effects of fossorial and herbivorous animals, help support an

unusually diverse ground flora (Hermann 1993). In the

northeast Florida-southwest Georgia arca, Small(1923)

recorded about 250 plant species in the longleaf-scrub oak

forests; the nearby Wade Tract supports more than 360

species (Tal1 Timbers Research Station herbarium list).

Longleaf pine ecosystems contain the greatest diversity of

plant life ever found in a temperate zone (Peet and Allard

1993), ofien with over 40 species per square meter and as

many as 140 species per 1000 m2. Of the 191 rare vascular

plant taxa associated with wiregrass in the Southeastern

United States, 122 are considered endangered or threatened

(Hardin and White 1989).

Pits and mounds associated with windthrown trees usually

do not persist because stumps are bumed and fiequent rains

have a leveling effect. Nonetheless, soil that erodes into old

pita, burrows, and stump cavems may be important in

maintaining heterogeneous substrates.

Important fungal diseases, such as brownspot needle blight

and root rot, are mitigated by fne. Multiple interactions have

been identified among pine trees, fungí, and key animal

species. The relationship between red heart decay in living,

old pines and the red-cockaded woodpecker, which

excavates cavities in decaying boles (Walters 199 1 ), is well

known. This bird’s selection of red heart trees that have been

suppressed and later released (Conner and O’Halloran 1987)

suggests that dominance-class diversity may be another

attribute of old growth. This certainly seemed to be the case

in the stand examined by Schwarz (1907). Weigl et al.

(1989) noted a three-way relationship among longleaf pine,
its mycorrhizae, and the southem fox squirrel, which eats

longleaf seeds and disperses mycorrhizae spores.

.Frequent tires inhibit soil microfauna and some

insectivorous small mammals that burrow underground

(Heyward and Tissot 1936). By contrast, fnes favor larger

herbivorous burrowers like gopher tortoises, pocket gophers,

and severa1 small rodents.

Wildlife diversity depends on a variety of habitat features

ofien formed by interacting disturbances. For example, at

least 145 vertebrate species have been identifled on the

Wade Tract. Important habitat components include large, old

pines and snags; younger age classes of pine; patchy forest

and groundcover; diverse ground flora; oak patches; open air

spaces; bare soil patches; fallen toarse woody debris; and

root-soil mounds tipped up by tree falls (Landers et al. 1990)

(see footnote 1). The red-cockaded woodpecker plays a key

role in that its cavities are used by many other birds and

mammals. R.T. Engstrom (see footnote 1) recorded the

woodpecker among 22 bird species that fiequent the longleaf

forest and are more abtmdant or found exclusively in old

growth. The long-term survival of the Coastal Plain fox

squirrel may also depend on mature to old-growth longleaf

(Kantola and Humphrey 1990). The gopher tortoise is a

primary grazer on xeric sites, and its burrows are used by at

least 60 vertebrate and 302 invertebrate species (Jackson

1989). The oak toad is also associated with the native,

grassy groundcover; and the flatwoods salamander depends

on large woody debris within grassy flatwoods (Motmt

1975).

Forest Dynamics and Ecosystem

Function

Functional and persistent old-growth longleaf pine

ecosystems depend on repeated fnes to prevent hardwood

domination, including Sres during the growing season,

which can slow or prevent recruitment of several fireresistant hardwoods into the midstory. Fire effects vary with

enviromnental gradients established by topography, soils,

and moisture levels. Sites intermediate in elevation and

moisture tend to burn more obten and more completely, but

less intensely, than either higher, xeric sites or lower, hydric

sites. Characteristic of the longleaf pines are mosaics of

longleaf pine-scrub oaks on sandhills grading to ahnost pure

longleaf pine on more mesic sites. Similar fire-maintained

associations that also span a range of sites fkom xeric to

hydric constitute habitats of South Florida slash pine.

Among the unique features of longleaf and South Florida

slash pine communities are the intricate relationships

between biological diversity and disturbance. Southeastem

pine uplands persisted for millennia tmder a regime of

frequent, widespread tires in a region of abundant rainfall. In

such a humid environment, community persistence was

probably reinforced by selection for both fire tolerance and

longevity in key species of pines, some oaks, and grasses

(Christensen 1981, Clewell 1986, Platt et al. 1988, Landers

199 1, Rebertus et al. 1993). The tire-tolerant grasses and

overstory with multiaged pine populations may be one of the

defining characteristics of old growth. This system’s

persistence is consistent with the concept of a true “climax,”

but only if lightning (an ignition source) is considered as

integral to climate, which determines the climax type

(Chapman 1932).

Fire plays an essential ecological role by clearing seedbeds

and recycling nutrients. Fire early in the growing season

promotes height growth in “grass stage” seedlings of

longleaf and South Florida slash pine, as well as wiregrass

seed production. It also synchronizes the flowering of many

herbaceous plants. Frequent fne tends to screen out broadleaved woody vegetation and favor a rich assortment of plant

life on the forest floor, including nitrogen-furing legumes

(mostly peremrials), low shrubs, and free-living soil

organisms.

Replacement of volatilized nutrients soon after a bum

contributes to a dynamic soil-nutrient equilibrium, as do the

nutrient- and water-absorbing functions of bunch grasses

(Clewell1986). Frequently burned pinelands have a humus

layer generated by grass roots that is less acidic with more

macronutrients available than that of unburned sites

(Wahlenberg 1946). Burning vegetation releases varying

quantities of nutrients into the atmosphere. It generally

increases the incorporation of phosphorus, potassium,

magnesium, calcium, nitrogen, and organic matter into

mineral soil, although short-term, fue-induced changes in the

proportion of elements will vary (Heyward and Bamette

1934, Christensen 1977, Hough 198 1, McKee 1982, Richter

et al. 1982, Gilliam 1988, Binkley et al. 1992). Long-term

research in Coastal Plain forests suggests that frequent

burning mobilizes calcium in the forest floor, retards soil

weathering processes, and maintains nutrient balances that

favor pine productivity (McKee 1982). Additional research

should center upon variation in nutrient flux, transformation,

and availability relative to geomorphic setting and fire

regime.

l l
Conservation and Management

Human alteration of pine forests, woodlands, and savannas

probably date back nearly 12,000 years. Historical records

suggest that Native Americans increased the fire frequency

in many areas, but they bumed in various seasons, adding to

rather than completely displacing natural fire regimes

(Robbins and Myers 1992, Frost 1993). Vast pinelands with

intact groundcover persisting through the era attest that

Indian agriculture did not greatly disrupt upland pine sites,

probably because of low site fertility.

By 1920, European settlement had brought agriculture,

livestock grazing, logging, and a nearly total loss of oldgrowth forests (Frost 1993). A 95percent reduction in fire

activity over the last 50 years (Simard and Main 1987), due

mainly to forest fire exclusion policies set in the 192O’s, has

also accelerated the loss of Ere-dependent ecosystems.

Longleaf pine, which was the dominant or principal tree

species on an estimated 92.5 million acres (37.5 million ha)

(Frost 1993), had dwindled to an estimated 3.8 million acres

(1.5 million ha) by 1985 (Kelly and Bechtold 1990). Since

then, its decline continued to about 3.2 million acres (1.3

million ha) by 1990 and to < 3 million acres ( 1.2 million ha)

by 1995 (Landers et al. 1995, Outcalt and Sheffield 1996) or

about 3 percent of the its presettlement area. Trees < 60

years old are few, and most of the remaining second growth

is 80 to 100 years old. However, < 2 percent of the Southeast

land area now contains longleaf pine, compared to 59

percent when settlement began (Frost 1993). Much of the

rest has been degraded by the absence of recurrent tire.

Conservation of naturally regenerated longleaf pine is a

challenge because two-thirds of the remaining acreage is

privately owned (Outcalt and Sheffield 1996), and much

private land is managed primarily for wood products. The

fnst step needed in a comprehensive conservation plan is to

inventory residual old growth, second growth that could

become old growth, and areas with native ground flora on

which pine could be restored. It would also be helpful to

identify sites with relatively undisturbed soils on which

native plant associations could be restored. Restoring and

maintaining fire-dependent ecosystems as a significant part

of southeastem landscapes would require regionwide

cooperation.

Successful restoration and maintenance of old growth will

generally require restoration of entire biological systems,

along with disturbance processes on which these systems

depend. A minimum for established plant communities

would be initial rough-reduction bums during the dormant

12

season, followed by a variable growing-season burn regime

(Robbins and Myers 1992). Procedures to reintroduce native

plants on cleared land, beginning with those species essential

to ecosystem processes, e.g., carbon storage, Ere ignition,

plant dispersal systems, food webs, nutrient and energy

cycling, and soil development, are needed.

Whether the pine species should be regarded primarily as

even-aged or tmeven-aged is an important question. It may

be that lar-ge, even-aged stands were most common near the

coas& particularly in major hurricane zones. Descriptions

suggest that, in many areas, mosaics of small, even-aged or

multiaged units persisted because of fire and localized

disturbances. In some areas, mosaics may also have resulted

from the gradual breakup, through attrition, of even-aged

stands established after a catastrophic event. Given the loss

of these ecosystems, old-growth diversity should be

recovered and maintained by developing a range of stand

structures and sizes. There is growing evidente that the

dynamic population structure of old-growth longleaf and

South Florida slash pine ecosystems is contingent upon

variation in the fiequency, behavior, and effects of fnes that

occur at different seasons (Platt et al. 1988, Doren et al.

1993). There is also a pressing need for guidelines suitable

for multiple-value forests. Research should focus on the

amount of vohune growth (perhaps 60 to 70 percent) that

could be harvested periodically without jeopardizing the

habitat of the flora and fauna indigenous to old-growth

ecosystems. Equally challenging is the maintenance of

understory plant communities through cycles of harvest and

regeneration.

Methods of forest management that would simulate oldgrowth conditions have natural analogs. Some of these

include frequent prescribed burning (especially in April to

June); harvesting in patches and keyed to major masting

events; partial cutting, such as single tree or group selection,

seed-tree reserve, or irregular shelterwood with varying

stand sizes; creating and maintaining a balanced distribution

of age and size, or both, and dominance classes by taking

advantage of or simulating natural disturbances; preparing

seedbeds by buming only; establishing seedlings through

natural regeneration or irregular plantings, if necessary;

minimizing disruption of soil and groundcover vegetation;

and leaving live cavity trees, snags, and downed logs. Lowintensity operations, e.g., hand planting, chainsaw felling,

and logging with draft animals or small equipment, may be

necessary to maintain ecological integrity within the context

of timber production.

Great challenges to sustain both ecological systems and

economic forestry operations in this region lie ahead.
Because so many forests have disappeared, the fate of pine

ecosystems now depends on the extent of restoration.

Solutions lie in developing networks of old-growth preserves

combined with managed forests dedicated to integrating

timber use and biodiversity, refmed stewardship schemes for

each objective, and economic incentives for landowners to

practice enviromnentally sensitive management.

The minimum area of such preserves will depend on the

elements to be perpetuated and the context in which the area

will be used. For example, the 206-acre (83-ha) Wade Tract

is a relatively fertile site within severa1 thousand acres of

second growth with much intact groundcover (Platt et al.

1988). Prescribed fnes maintain the natural plant community

there during the growing season. On the other hand, the

Wade Tract contains only three clusters of cavity trees used

by the red-cockaded woodpecker, a species that requires old

pines for nesting and extensive pine stands for foraging.

Considering estimates for a breeding core of 3 10 to 390

pairs to sustain a viable population and a minimum territory

of 200 acres (8 1 ha) per pair, the minimum area of

acceptable habitat required may be 80,000 acres (32,374 ha)

or more (Southeast Negotiation Network 1990). However,

periodic genetic information may reduce the minimum size

of a breeding core and the acreage required for a stable

population (Walters 199 1).

Because of the uncertainty of area parameters, it may be

useful to consider a minimum building block for stewardship

of old growth within a younger forest. For example, 125

acres (5 1 ha) of old growth should generally encompass a

mosaic of pine age classes, the minimum territory for a

single red-cockaded woodpecker clan, the range for a

breeding group of fox squirrels (Weigl et al. 1989), and

space for many other animals and plants common to old

growth. However, the critica1 area necessary for populations

of most species and the importance of community

interactions are not yet known. Viability of the ecosystem

will likely increase with the size and number of old-growth

reserves covering the principal sites and community types,

shorter distances between reserves, and greater similarities

in composition and structure among the reserves and the

areas separating them. Long-term research will be needed to

determine minimm areas required for long-term

maintenance of complete old-growth systems.

Representative Old-Growth Stands

For the remaining old-growth areas, landowners have

controlled access. Few sites have been frequently burned, so

most tracts do not entirely represent natural community

structure and composition. Fire must be reintroduced to

restore plant communities, including natural pine

regeneration. Ecological research and restoration, or both,

and stewardship are being planned or implemented for al1

tracts listed below.

Longleaf Pine

l Boyd Tract-Weymouth Woods Sandhills Nature Preserve,

Moore Cotmty, North Carolina

l Big Woodsareenwood Plantation, Thomas County,

Georgia

l Hauss Park-Flomaton, Escambia County, Alabama

(Champion Intemational Corp.)

l Patterson Tract-Okaloosa County, Florida (Eglin Air

Forte Base)

l Wade Tract-Thomas Cotmty, Georgia (Tal1 Timbers, Inc.)

South Florida Slash Pine

l Lostman’s Pines-Everglades National Park, Florida

l Raccoon Point-Big Cypress National Preserve, Florida

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15

Landers, J. Larry; Boyer, William D. 1999. An old-growth defínition for upland

longleaf and south Florida slash pine forests, woodlands, and savannas. Gen. Tech.

Rep. SRS-29. Asheville, NC: U.S. Department of Agriculture, Forest Service,

Southem Research Station. 15 p.

Upland longleaf pine forests, woodlands, and savannas once occupied most of the

Atlantic and Gulf Coastal Plains t?om southeastem Virginia south through the

northem two-thirds of Florida and west to east Texas, with extensions into the

Piedmont and mountains of Alabama and northwest Georgia. South Florida slash pine

is native to the southem half of peninsular Florida. The probable stmcture of and

variability in old-growth forests, woodlands, and savannas of these two overlapping

and fire-dependent ecosystems are described based on al1 available information. A

number of old-growth attributes are listed for both species, including estimates of the

variability that can be expected in these attributes.

Keywords: Longleaf pine, old growth, savannas, South Florida slash pine,

Southeastem United States, woodlands.

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