United States

Department of

Agriculture

Forest Service

Southern 

Research Station

General Technical

Report SRS-83

Restoration of 

Longleaf Pine Ecosystems 

Dale G. Brockway, Kenneth W. Outcalt, 

Donald J. Tomczak, and Everett E. Johnson
Dale G. Brockway, Research Ecologist, USDA Forest Service, Southern 

Research Station, Auburn, AL 36849; Kenneth W. Outcalt, Research 

Plant Ecologist, USDA Forest Service, Southern Research Station, Athens, 

GA 30602; Donald J. Tomczak, Silviculturist, USDA Forest Service, 

Southern Region, Atlanta, GA 30367; and Everett E. Johnson, Director, 

Solon Dixon Forestry Education Center, School of Forestry and Wildlife 

Sciences, Auburn University, Andalusia, AL 36420.

The Authors

July 2005

Southern Research Station

P.O. Box 2680

Asheville, NC 28802

The use of trade or firm names in this publication is for reader information and does not imply endorsement 

of any product or service by the U.S. Department of Agriculture or other organizations represented here.

PESTICIDE PRECAUTIONARY STATEMENT

This publication reports research involving pesticides. It does not contain recommendations for their use, 

nor does it imply that the uses discussed here have been registered. All uses of pesticides must be registered 

by appropriate State and/or Federal agencies before they can be recommended.

CAUTION: Pesticides can be injurious to humans, domestic animals, desirable plants, and fish or other 

wildlife if they are not handled or applied properly. Use all pesticides selectively and carefully. Follow 

recommended practices for the disposal of surplus pesticides and their containers.
i

Restoration of 

Longleaf Pine Ecosystems

Dale G. Brockway, Kenneth W. Outcalt, 

Donald J. Tomczak, and Everett E. Johnson
ii
iii

Contents

Southern Forest Environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Longleaf Pine Ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

 Longleaf Pine Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

 Ecological Significance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

History of Longleaf Pine Ecosystems. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Social, Political, and Economic Context . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

Restoration Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

 Defining Restoration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

 Measuring Success . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

 Future Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

Restoration Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

 Restoration Framework . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

 Selecting Techniques . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

 Restoration Prescriptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

 Restoration Efforts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

Costs and Benefits Associated with Restoration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

 Estimating Restoration Costs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

 Benefits of Restored Longleaf Pine Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Influence of Forestland Ownership . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

 Regional Patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

 Land Ownership Patterns in Longleaf Pine Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

 Influence of Ownership on Restoration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

Research and Management Needs and Policy Implications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

 Scientific Knowledge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

 Management Information . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

 Policy Implications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
iv
1

Restoration of Longleaf Pine Ecosystems

Dale G. Brockway, Kenneth W. Outcalt, Donald J. Tomczak, and

Everett E. Johnson

Abstract

Longleaf pine (Pinus palustris) ecosystems once occupied 38 million ha 

in the Southeastern United States, occurring as forests, woodlands, and 

savannas on a variety of sites ranging from wet flatwoods to xeric sandhills 

and rocky mountainous ridges. Characterized by an open parklike structure, 

longleaf pine ecosystems are a product of frequent fires, facilitated by the 

presence of fallen pine needles and bunchgrasses in the understory. Timber 

harvest, land conversion to agricultural and other nonforest uses, and alteration of fire regimes greatly reduced longleaf pine ecosystems, until only 

1.2 million ha remained in 1995. Longleaf pine ecosystems are among the 

most species-rich ecosystems outside the tropics. However, habitat loss 

and degradation have caused increased rarity of many obligate species. 

The lack of frequent surface fires and the proliferation of woody plants in 

the understory and midstory have greatly increased the risk of additional 

longleaf pine ecosystem losses from catastrophic fire.

Because longleaf pine still exists in numerous small fragments throughout 

its range, it is reasonable to conclude that it can be restored. Restoration 

efforts now underway use physical, chemical, and pyric methods to reestablish the natural structure and function in these ecosystems by adjusting 

species composition, modifying stand structure, and facilitating ecological 

processes, such as periodic fire and longleaf pine regeneration. The ecological, economic, and social benefits of restoring longleaf pine ecosystems 

include (1) expanding the habitat available to aid in the recovery of numerous imperiled species, (2) improving habitat quality for many wildlife 

species, (3) producing greater amounts of high-quality longleaf pine timber 

products, (4) increasing the production of pine straw, (5) providing new 

recreational opportunities, (6) preserving natural and cultural legacies, and 

(7) creating a broader range of management options for future generations.

Keywords: Biological diversity, bluestem grasses, disturbance, fire ecology, 

gopher tortoise, Pinus palustris Mill., red-cockaded woodpecker, wiregrass. 

Southern Forest Environment

Longleaf pine ecosystems occur in the southern forest region

of the 13 States in the Southeastern United States. This region

is bounded by the Atlantic Ocean to the east, the Gulf of 

Mexico to the south, the Great Plains grasslands to the west, 

and the interior plateaus, highlands, and mountains to the 

north (Miller and Robinson 1995). Lower elevations in this 

region consist of broad Coastal Plains that extend along the 

shores of the Atlantic Ocean and Gulf of Mexico. In the 

western part of the region, the Coastal Plain is interrupted by 

the Mississippi River’s alluvial plain. Middle elevations, in 

the eastern portion of the region, are occupied by the Piedmont, a broad area of uplands which parallels the Atlantic 

Coastal Plain. Numerous sandhills occur along the contact 

between the Coastal Plains and the Piedmont, as well as along

the Lake Wales Ridge of central Florida and beach ridges in

northwestern Florida. Upper elevations, extending to nearly

2000 m, are dominated by the Blue Ridge Mountains, Cumberland Plateau, and Ouachita and Ozark Mountains. Longleaf 

pine forests naturally occur at elevations < 600 m (with most 

occurring at < 200 m) in 9 of the States in this region (all 

those except Oklahoma, Arkansas, Tennessee, and Kentucky, 

where cooler environments predominate) (Boyer 1990b).

Except in the highlands, most of the southern region is 

characterized by a humid subtropical climate (Bailey 1995). 

Temperatures are generally high, with minimums during 

January ranging from 0 °C to 13 °C and maximums during 

July ranging from 29 °C to > 35 °C. Annual precipitation 

varies from 1040 to 1750 mm and is well distributed through 

the year. During the late summer and fall seasons, hurricanes 

developing over the Atlantic Ocean move westward and 

frequently affect Coastal Plain forests in this region. Such 

tropical storms are one of the principal large-scale damaging 

agents for longleaf pine forests growing near the seacoast. 

Growing seasons are comparatively long in the southern 

region, with those near upper elevations between 160 and 

220 days and those in the lower Coastal Plains of central 

Florida > 300 days. 

In the highlands, soils are derived from parent materials 

largely consisting of granite, quartzite, schist, slate, sandstone, shale, limestone, and dolomite. At lower elevations, 

parent materials are generally marine sediments ranging from

deep, coarse, excessively drained sands to poorly drained 

clays. Surface soils derived from these progenitors are generally sandy, acidic, low in organic matter, and relatively 

infertile (Boyer 1990b). The predominant forest soils in the 

southern region (outside peninsular Florida) are the Ultisols. 

The red-yellow color in the profile of these soils results from

the translocation of iron and aluminum from a lighter-colored

upper horizon to a darker red B horizon below (Pritchett 

1979). Typic Paleudults and Plinthic Paleudults are the 

Ultisols most commonly associated with longleaf pine 

forests. Entisols and Spodosols are the other soil orders most 

frequently observed to support longleaf pine. Entisols are 

deep sands, with very weak horizon development, found 
2

on relatively xeric sandhills. Most commonly occurring as 

Quartzisamments, these soils range from elevations of 3 m 

in coastal Florida to 185 m in the sandhills of the Carolinas, 

Georgia, northwestern Florida, and the central ridge of 

Florida (Boyer 1990b). Spodosols, principally Aquods, are 

characteristic of lower Coastal Plain flatwoods, which have 

wet sandy soils with a shallow water table that is at or near 

the ground surface during the rainy season.

Southern forests occupy 87 million ha of a vast and diverse 

landscape which supports a great number of ecosystems and 

species, including > 60 forest types (Walker and Oswald 

2000). The forests of this region have great commercial 

importance and produce ~ 16 percent of the world’s roundwood supply, more than any other individual nation. The 

area of naturally regenerated pine forests in the South has

declined as a result of natural pine harvesting and establishment of plantations, succession to hardwoods, and conversion

to nonforest uses (i.e., urban development or agriculture). 

The area occupied by pine plantations in the region is forecast to increase by 67 percent to 22 million ha by the middle 

of the 21st century (Prestemon and Abt 2002). The influence 

this change will have (along with other stresses) on rare 

forest communities is uncertain. However, it is widely agreed

that habitat reduction and loss are the principal causes of 

increasing species endangerment throughout the region. 

Longleaf pine forests are recognized as being among the 

most endangered of southern ecosystems (Wear and Greis 

2002).

Highlands are occupied by various forest types including 

Fraser fir (Abies fraseri) and red spruce (Picea rubens) mixed

with hardwoods, oak-hickory (Quercus-Carya), table mountain pine (Pinus pungens), shortleaf pine (P. echinata), loblolly pine (P. taeda), Virginia pine (P. virginiana), and eastern

redcedar (Juniperus virginiana) (Walker and Oswald 2000). 

Small amounts of montane longleaf pine, in generally poor 

condition, still occur in northern Alabama and Georgia 

(Boyer 1990b). However, most natural stands of longleaf pine

in the mountain province have declined greatly, as timber 

harvesting and fire exclusion resulted in their replacement by

loblolly pine, shortleaf pine, and hardwoods. The Piedmont 

is largely a mixture of southern pines and hardwoods, supporting loblolly pine and shortleaf pine plantations, oakhickory forests, oak-pine forests, and stands of Virginia pine

(Walker and Oswald 2000). The province also supports longleaf pine on xeric sands and mesic uplands, and riparian 

forests along stream bottoms (Golden 1979). The once 

extensive longleaf pine forests of the Piedmont have largely 

been converted to loblolly pine plantations through timber 

harvesting and replanting (Schultz 1997) or replaced by 

hardwoods that have flourished in the absence of frequent 

fire. The Coastal Plain is characterized by forests of loblolly 

pine and shortleaf pine, longleaf pine and slash pine (P. 

elliottii), oak-pine, upland hardwoods, sand pine (P. clausa) 

and pond pine (P. serotina), Virginia pine, eastern redcedar, 

Atlantic white-cedar (Chamaecyparis thyoides), baldcypress 

(Taxodium distichum), and bottomland hardwoods (Walker 

and Oswald 2000). On the lowest terraces along the coast 

(at elevations < 8 m), poorly drained but not permanently 

flooded soils occur over extensive flatwoods areas, which are

commonly occupied by pines and hardwoods. Although 

formerly dominated by longleaf pine, many of these flatwoods sites have been converted to slash pine and loblolly 

pine plantations. The uplands of the middle and upper Coastal

Plains are typically well-drained sites supporting forests of

southern pines and hardwoods. While longleaf pine also 

once dominated these landscapes, it has been largely replaced

by plantations of loblolly pine (Schultz 1997). The xeric 

sandhills located in Florida and along the fall line between 

the Piedmont and the Coastal Plain are still largely occupied 

by longleaf pine and its scrub oak associates. The primary 

threat to the remaining longleaf pine is the absence of 

frequent fire, which results in encroachment by sand pine, 

hardwoods, and understory scrub species. 

Longleaf Pine Ecology

Longleaf Pine Ecosystems

Longleaf pine forests were once among the most extensive 

ecosystems in North America (Landers and others 1995). 

Prior to European settlement, these forests occupied ~ 38 

million ha in what is now the Southeastern United States 

(Frost 1993). Travelers in this region during the late 18th and 

early 19th centuries reported that there were vast areas in 

which longleaf pine covered > 90 percent of the landscape 

(Bartram 1791, Williams 1837). The native range of longleaf 

pine (fig. 1) extends along the Gulf and Atlantic Coastal 

Plains from Texas to Virginia, and well into central Florida 

and the Piedmont and mountains of northern Alabama and 

northwest Georgia (Boyer 1990b, Stout and Marion 1993). 

Longleaf pine occurs in forests, woodlands, and savannas on 

sites that range from wet, poorly drained flatwoods to mesic 

uplands, xeric sandhills, and rocky mountainous ridges 

(Boyer 1990b). Associated species are often influenced by 

site quality, but longleaf pine is a species of great ecological 

amplitude, and its occurrence is largely unaffected by the 

soil type (Gilliam and others 1993, Kalisz and Stone 1984), 

except that there is greater early competition on sites of 

higher quality. 
3

An open, parklike stand structure (fig. 2) is a distinguishing 

characteristic of longleaf pine ecosystems (Edmisten 1963, 

Schwarz 1907, Wahlenberg 1946). Naturally occurring 

longleaf pine forests are typically an uneven-aged mosaic of 

even-aged patches distributed across the landscape, which 

vary in size, shape, structure, composition, and density, and

contain numerous embedded special habitats such as stream 

bottoms, wetlands, and seeps (Brockway and Outcalt 1998,

Hilton 1999, Platt and Rathbun 1993). The natural variability

of these ecosystems makes them excellent habitat for game 

animals such as white-tailed deer (Odocoileus virginianus), 

wild turkey (Meleagris gallapavo), and northern bobwhite 

(Colinus virginianus) and numerous nongame and rare animal

species, including fox squirrel (Sciurus niger), southeastern 

pocket gopher (Geomys pinetis), Bachman’s sparrow (Aimophila aestivalis), brown-headed nuthatch (Sitta pusilla), redcockaded woodpecker (Picoides borealis), gopher tortoise 

(Gopherus polyphemus), eastern indigo snake (Drymarchon 

corais couperi), eastern diamondback rattlesnake (Crotalus 

adamanteus), and flatwoods salamander (Ambystoma cingulatum) (Brockway and Lewis 2003, Crofton 2001, Engstrom 

1993, Guyer and Bailey 1993, Kantola and Humphrey 1990). 

In the west Gulf Coastal Plain, longleaf pine understories are

dominated by bluestem grasses (Schizachyrium scoparium

and Andropogon spp.). From Florida north and eastward, 

longleaf pine is typically associated with wiregrass (Aristida

stricta, also known as pineland threeawn, and A. beyrichiana,

Figure 1—Native range of longleaf pine and physiographic provinces of the Southeastern United States (Little 1971, Miller and Robinson 1995).
4

Beyrich threeawn). Because fallen pine needles and understory grasses facilitate the ignition and spread of fire during 

the growing season, these plant communities often contain 

relatively few shrubs or hardwood trees (Landers 1991). 

Although such woody plants may be more numerous on 

mesic sites, their stature is typically limited by frequent 

burning. Quercus, Ilex, and Serenoa are common tree and 

shrub associates of longleaf pine at various locations within 

its native range. Longleaf pine ecosystems support a great 

variety of herbaceous plant species (table 1). The high 

diversity of understory plants per unit area makes these 

ecosystems among the most species-rich plant communities 

outside the tropics (Peet and Allard 1993). 

Throughout its domain, longleaf pine is closely associated 

with frequent surface fires (Brockway and Lewis 1997, 

Garren 1943, Outcalt 2000, Wright and Bailey 1982). In 

these ecosystems, longleaf pine and bunchgrasses function 

together as keystone species that facilitate but are resistant to 

fire (Noss 1989, Platt and others 1988b). They are relatively 

long-lived and retain nutrients and water to a degree that 

reinforces their site dominance and minimizes change in 

the plant community following disturbance (Landers and 

others 1995). Fire can spread very quickly through a finefuel matrix composed of highly flammable longleaf pine 

needles and the leaves of several bunchgrasses (Abrahamson 

and Harnett 1990, Landers 1991). The benefits of periodic 

fire include (1) maintaining the physiognomic character of 

longleaf pine bunchgrass ecosystems by excluding invasive 

plants that are poorly adapted to fire, (2) preparing a seedbed 

favorable for the regeneration of longleaf pine seedlings, 

(3) reducing the density of understory vegetation and thus 

providing microsites for a variety of herbaceous plants,

(4) stimulating increased seed production by native grasses,

(5) releasing nutrients immobilized in accumulated phytomass for recycling to the infertile soil and subsequently more

Figure 2—Longleaf pine–wiregrass ecosystem on xeric sandhill site in Florida.
5

Table 1—Plants associated with longleaf pine forest ecosystems

Category and common name Scientific name

Trees

 Pignut hickory Carya glabra 

 Sweetgum Liquidambar styraciflua 

 Shortleaf pine Pinus echinata

 Slash pine P. elliottii

 Longleaf pine P. palustris

 Southern red oak Quercus falcata

 Bluejack oak Q. incana

 Turkey oak Q. laevis

 Laurel oak Q. hemisphaerica

 Blackjack oak Q. marilandica

 Water oak Q. nigra

 Post oak Q. stellata

Shrubs 

 Polecat bush Asimina incana

 Pawpaw A. triloba

 Groundsel tree Baccharis halimifolia

 American beautyberry Callicarpa americana

 Rosemary Ceratiola ericoides

 Coastal sweet pepperbush Clethra alnifolia

 Garberia Garberia fruticosa

 Dwarf huckleberry Gaylussacia dumosa

 Blue huckleberry G. frondosa

 Gallberry Ilex glabra

 Yaupon I. vomitoria

 Gopher apple Licania michauxii

 Rusty staggerbush Lyonia ferruginea

 Coastalplain staggerbush L. fruticosa

 Fetterbush lyonia L. lucida

 Wax myrtle Myrica cerifera

 Prickly pear Opuntia humifusa

 Chokeberry Pyrus arbutifolia

 Dwarf live oak Q. minima

 Running oak Q. pumila

 Winged sumac Rhus copallinum

 Scrub palmetto Sabal etonia

 Saw palmetto Serenoa repens

 Sparkleberry Vaccinium arboreum

 Shiny blueberry V. myrsinites

 Deerberry V. stamineum

 Adam’s needle Yucca filamentosa

 Coontie Zamia pumila

Vines 

 Yellow jessamine Gelsemium sempervirens

 Morning glory Ipomoea spp.

 Virginia creeper Parthenocissus quinquefolia

Category and common name Scientific name

Vines (cont.) 

 Blackberry, dewberry, Rubus spp.

 raspberry

 Greenbrier Smilax spp.

 Poison ivy Toxicodendron radicans

 Grape Vitis spp.

Graminoids

 Splitbeard bluestem Andropogon ternarius 

 Broomsedge bluestem A. virginicus

 Beyrich threeawn, wiregrass Aristida beyrichiana

 Pineland threeawn, wiregrass A. stricta

 Switchcane Arundinaria gigantea

 Common carpetgrass Axonopus fissifolius

 Ware’s hairsedge Bulbostylis warei

 Sedge Carex spp.

 Toothache grass Ctenium aromaticum

 Nutsedge Cyperus spp.

 Rosette grass Dichanthelium spp.

 Purple lovegrass Eragrostis spectabilis

 Skeletongrass Gymnopogon spp.

 Cutover muhly Muhlenbergia capillaris

 Panicgrass Dichanthelium spp.

 Paspalum Paspalum spp.

 Beakrush Rhynchospora spp.

 Sugarcane Saccharum spp.

 Little bluestem Schizachyrium scoparium

 Nutrush Scleria spp.

 Lopsided indiangrass Sorghastrum secundum

 Curtis’ dropseed Sporobolus curtissii

 Pineywoods dropseed S. junceus

 Purpletop tridens Tridens flavus

 Sand grass Triplasis spp.

Forbs

 Figwort Scrophularia spp.

 Milkweed Asclepias spp.

 Scaleleaf aster Symphyotrichum adnatum

 Bushy aster A. dumosus

 Stiff-leaved aster A. linariifolius

 White-topped aster A. paternus

 Pinebarren white-topped aster Oclemena reticulata

 Wild indigo Baptisia lanceolata

 Soft greeneyes Berlandiera pumila

 Florida lady’s nightcap Bonamia grandiflora

 Vanillaleaf, deer tongue Carphephorus 

 odoratissimus

 Partridge pea Cassia chamaecrista

continued
6

rapid uptake by plants, (6) improving forage for grazing,

(7) enhancing wildlife habitat, (8) controlling harmful insects

and pathogens, and (9) reducing fuel levels and the wildfire 

hazard (Brennan and Hermann 1994, Dickmann 1993, 

Haywood and others 2001, Landers and Boyer 1999, Lemon 

1949, McKee 1982, Outcalt 1994, Wade and Lewis 1987, 

Wade and Lundsford 1990). Fire at intervals as frequent as 

2 to 4 years may provide such benefits in these ecosystems 

without need for measures to protect regeneration. However, 

recurrent fire in any season can be responsible for growth 

losses in overstory longleaf pines ranging from 10 to 18 

percent (Boyer 2000). Trading these losses for the benefits 

described above seems worthwhile, especially because the 

negative effect of fire on growth of longleaf pine trees is 

greatest at younger ages and diminishes with time.

Prior to landscape fragmentation, natural fires occurred every

2 to 8 years throughout much of the region (Abrahamson and

Harnett 1990, Christensen 1981). Longleaf pine was dominant over large areas primarily because it is more tolerant 

of frequent fire than competing species with thinner-barked 

seedlings. Although longleaf pine seedlings are very susceptible to fire-caused mortality during the first year following 

germination, they become increasingly resistant to fire in 

subsequent years. A unique adaptation of longleaf pine to its

fire-prone environment is a seedling “grass stage,” during 

which root growth is favored and the seedling top remains 

a tuft of needles surrounding a large terminal bud. Because 

there is no stem, no cambium is directly exposed to damage 

from surface fires. When sufficient root reserves have accumulated, grass-stage longleaf pine seedlings “bolt” or “rocket,” 

rapidly growing 1 to 2 m in a short time. Such rapid growth 

puts the terminal bud beyond the lethal reach of most surface

fires. Larger longleaf pine trees have relatively thick bark 

that protects cambial tissue from the lethal heating of surface

fires (Wahlenberg 1946). Fires assist in the natural pruning 

of longleaf pine, creating a clear bole between the crown 

and any accumulated surface fuels. Surface fires are thereby 

prevented from easily moving into the canopy. The tendency 

of longleaf pine to regenerate more successfully in forest 

Table 1—Plants associated with longleaf pine forest ecosystems (cont.)

Category and common name Scientific name

Forbs (cont.)

 Maryland goldenaster Chrysopsis mariana

 Atlantic pigeonwings Clitoria mariana

 Treadsoftly Cnidoscolus stimulosus

 Rabbitbells Crotalaria rotundifolia

 Healing croton Croton argyranthemus

 Gulf croton, beachtea C. punctatus

 Ticktrefoil Desmodium spp.

 Elephant’s foot Elephantopus tomentosus

 Dogtongue buckwheat Eriogonum tomentosum

 Dogfennel Eupatorium capillifolium

 Downy milkpea Galactia volubilis

 Coastal bedstraw Galium hispidulum

 Sunflower Helianthus spp.

 Hawkweed Hieracium gronovii

 St. Johns-wort Hypericum spp.

 Bush clover Lespedeza spp.

 Blazing star Liatris spp.

 Carolina lily Lilium michauxii

 Shortleaf lobelia Lobelia brevifolia

 Sensitive plant Mimosa spp.

 Partridgeberry Mitchella repens

 Woodsorrel Oxalis spp.

 Virginia plantain Plantago virginica

 Orange milkwort Polygala lutea

Category and common name Scientific name

Forbs (cont.)

 Candyroot P. nana

 Wand blackroot Pterocaulon virgatum

 Meadowbeauty Rhexia spp.

 Dollarleaf Rhynchosia reniformis

 Blackeyed susan Rudbeckia hirta

 Helmet flower Scutellaria integrifolia

 Seymaria Seymaria spp.

 Rosinweed Silphium spp.

 Jeweled blue-eyed grass Sisyrinchium solstitiale

 Queen’s-delight Stillingia sylvatica

 Sidebeak pencilflower Stylosanthes biflora

 Hoarypea Tephrosia spp.

 Noseburn Tragia spp.

 Forked blue curls Trichostema dichotomum

 Ironweed Veronica angustifolia

 Violet Viola spp.

 Carolina yelloweyed grass Xyris caroliniana

Ferns

 Cinnamon fern Osmunda cinnamomea

 Western brackenfern Pteridium aquilinum

Lichens

 Reindeer lichen Cladonia subtenuis
7

openings than directly beneath mature trees where long-term 

survival is poor (Brockway and Outcalt 1998) limits the 

development of ladder fuels near the crowns of adult trees.

Longleaf pines have the biological potential to live for 500

years, but they seldom survive this long because they exist 

in an environment that is subject to frequent disturbance 

(Engstrom and others 2001, Palik and Pederson 1996). 

Damaging tropical storms, such as hurricanes and the tornadoes associated with them, may fell extensive areas of trees

and open numerous gaps in the canopy of longleaf pine 

forests (Croker 1987). Lightning is another important mortality agent, typically killing individual trees but sometimes 

striking small groups of trees (Komarek 1968, Palik and 

Pederson 1996, Taylor 1974). Although insect infestations 

are uncommon, annosus root rot (Heterobasidion annosum), 

pitch canker (Fusarium moniliforme var. subglutinans), cone 

rust (Cronartium strobilinum), and other pathogens may 

infect longleaf pine (Boyer 1990b). Epidemics of brown-spot

disease (Mycosphaerella dearnessii) occasionally occur in

young longleaf pines, and this pathogen is usually fatal unless

a surface fire consumes the infected needles and cleanses the

stand of innoculum (Boyer 1990b, Wright and Bailey 1982).

Fire acts as a principal disturbance agent, influencing many

attributes and processes in longleaf pine ecosystems 

(Christensen 1981, Noss 1989). The long-term result of frequent surface fires is a forest composed primarily of pyrophytic vegetation and other plant and animal species whose

life cycles are adapted to the prevailing disturbance regime 

(Brockway and Lewis 1997, Engstrom and others 2001, 

Landers 1991). The structure, pattern, and biological diversity in these forest ecosystems are maintained by a combination of disturbance events and site factors. Variation in 

lightning strikes, tree mortality, and animal interactions at 

local scales, and wind storms, soil attributes, and hydrologic 

regimes at broader scales influence the landscape mosaic. 

Such forces acting across site gradients provide for large 

living trees, snags, coarse woody debris, hardwood thickets, 

and forest canopy gaps in this disturbance-prone, yet largely 

stable ecosystem. 

Longleaf pine is shade-intolerant, and its regeneration is

largely confined to canopy gaps (Wahlenberg 1946). Establishment of pine seedlings in gaps at different times results in

a network of forest patches at various stages of development 

dispersed across the landscape (Pickett and White 1985). 

These gap-phase regeneration dynamics give rise to the pattern of even-aged patches distributed across an uneven-aged 

landscape mosaic commonly observed in natural longleaf 

pine ecosystems (Palik and others 1997). Canopy gaps resulting from a variety of disturbance agents have recently been

recognized as ecologically important features driving the

forest cycle through open, growth, and closed phases (Coates

and Burton 1997, Whitmore 1989) and may serve in longleaf

pine forests as vehicles through which to apply silviculture 

that is modeled after natural disturbance (Palik and others 

2002). 

Conditions that ensure reliable regeneration are essential for

restoring and sustaining longleaf pine forest ecosystems and

can be achieved by combining frequent fire with natural 

regeneration techniques (Boyer and White 1990). Both abundance and growth of seedlings are negatively related to the 

presence of adult longleaf pines, which have a competitive 

influence on seedlings that extends > 15 m into forest gaps

(Boyer 1963; Smith 1955; Walker and Davis 1954, 1956). 

Naturally regenerated longleaf pine seedlings on xeric 

sandhills cluster near the center of canopy gaps, in an area

corresponding to a fine-root gap surrounded by a seedling 

exclusionary zone (SEZ) approximately 12 to 16 m wide

(Brockway and Outcalt 1998). Over many years of competition and repeated fires, seedling density declines more rapidly

in the SEZ than near the gap center. This fine-root gap and

SEZ structure has not been observed during the short term in 

newly created gaps on more mesic sites containing greater

numbers of woody plants (Gagnon and others 2003, McGuire

and others 2001, Palik and others 1997). While a number of

even-aged and uneven-aged approaches appear to be compatible with restoring and sustaining longleaf pine ecosystems

(Boyer 1993, Farrar 1996, Guldin 1996), silvicultural methods

such as group selection and irregular shelterwood, which 

mimic the pattern of natural disturbance and thus facilitate 

the normal ecological process of forest regeneration, are 

especially well suited for these purposes (Brockway and 

Outcalt 1998).

Ecological Significance

The complex natural pattern and disturbance-mediated processes that characterize longleaf pine forests create extraordinarily high levels of biological diversity in these ecosystems. 

The great number of plant species per unit area qualifies longleaf pine forests as among the most species-rich terrestrial 

ecosystems in the temperate zone. As many as 140 species 

of vascular plants have been observed in a 1000-m2 area, 

and > 40 species/m2 have been recorded in many longleaf 

pine communities (Peet and Allard 1993). A large number 

of these plant species are restricted to or found principally 

in longleaf pine habitats. Not surprisingly, many animal 

species also depend on longleaf pine ecosystems for much 

of their habitat. Among these are two increasingly rare 

animals that function as important primary excavators. Tree 

cavities created by red-cockaded woodpeckers and ground 

burrows dug by gopher tortoises provide homes for a wide 

variety of secondary users such as insects, snakes, birds, 
8

History of Longleaf Pine Ecosystems

During the Wisconsinan Ice Age (~ 40,000 to 12,000 years

BP), forests in the southern region consisted of boreal elements (Picea, Pinus) and temperate species (Carya, Castanea,

Ostrya, Quercus) intermixed in a pattern that varied both 

spatially and temporally with the ebb and flow of the vast ice

sheet farther to the north (Delcourt 1980, Watts 1970, Watts 

and others 1992). As the continental glacier retreated, southern forests became dominated by oaks and a diverse array 

of deciduous hardwoods after 12,000 years BP (Watts 1971, 

Watts and Hansen 1988, Watts and others 1992). Longleaf 

pine, moving northward and eastward from its Ice Age refuge

in southern Texas or northern Mexico (Schmidtling and 

Hipkins 1998), became established in the lower Coastal 

Plain ~ 8,000 years ago (Watts and others 1992) and during 

the ensuing 4,000 years spread throughout the Southeast 

(Delcourt and Delcourt 1987). This several-thousand-year 

pre-Columbian time period coincides with the interval 

during which populations of Native Americans flourished 

throughout the region, and their use of fire is thought to be 

related to the development and maintenance of longleaf pine 

forest ecosystems (Landers and Boyer 1999, Pyne 1997, 

Schwartz 1994).

Native Americans frequently used fire to manipulate their 

environment (Anderson 1996, Carroll and others 2002, 

Robbins and Myers 1992, Stanturf and others 2002), and 

early European settlers adopted the practice of periodically 

burning nearby forests and woodlands to improve forage 

quality for cattle grazing and discourage the encroachment 

of shrubby undergrowth. Although well adapted to frequent 

surface fires, longleaf pine was not well adapted to other 

disturbances brought by European settlement. As a result of 

the cumulative impacts brought by three centuries of changing land use, longleaf pine forests declined dramatically. By 

1900, logging, harvest of naval stores (chemicals derived 

from pine resin), and agriculture had reduced by more than 

half the area dominated by longleaf pine (Frost 1993). Logging continued until only fragments of the original longleaf 

pine forest remained in 1935. Second-growth longleaf pine 

stands became reestablished on only one-third of the sites 

previously occupied (Wahlenberg 1946). Harvest of these 

second-growth stands, often followed by conversion to other 

southern pines or urban development, continued through 

1985 (Kelly and Bechtold 1990). Over the next decade, longleaf pine was further reduced until it occupied < 5 percent of 

its original area (Outcalt and Sheffield 1996). 

The effects of settlement on longleaf pine forests were 

initially minor, with harvesting limited to areas near early 

towns and villages where log structures were constructed 

(Croker 1987). Later, lumber was cut from longleaf pine 

and mammals (Engstrom 2001, Jackson and Milstrey 1989, 

Speake 1981).

The longleaf pine forests and savannas of the Southeastern 

Coastal Plain are among the most critically endangered 

ecosystems in the United States, now occupying < 3 percent 

of their original extent (Noss and others 1995, Ware and 

others 1993). Extreme habitat reduction is the primary cause

for increasing rarity of 191 taxa of vascular plants and several

terrestrial vertebrate species that are endemic to or exist 

largely in longleaf pine communities (Hardin and White 

1989, Walker 1993). Habitat loss has principally resulted 

from conversion of longleaf pine forests to other land uses 

(i.e., agriculture, industrial pine plantations, and urban development), landscape fragmentation, and interruption of natural

fire regimes (Landers and others 1995, Wear and Greis 2002).

Edaphic variation and land use history, particularly as related

to soil disturbance or agriculture, have significant influences 

on the diversity of understory plants (Hedman and others 

2000, Rodgers and Provencher 1999). Frequent fire in differing seasons is appropriate for maintaining a greater variety 

of native plant species (Hiers and others 2000, Kirkman and

others 1998a). However, long-term exclusion of fire typically

results in depressed species diversity, development of a substantial hardwood understory and midstory, and accumulation of a thick layer of forest litter (Brockway and Lewis 

1997, Kush and Meldahl 2000). Such buildup of forest fuel 

can transform surface fires into crown fires that can have 

catastrophic effects on the rare plants and animals. Safe and 

effective reintroduction of fire into long-unburned forests 

remains the critical conservation challenge (Wear and Greis 

2002). 

Longleaf pine bunchgrass ecosystems are also vital to the 

maintenance of many biotic communities embedded in the

forest landscape matrix (Landers and others 1990). Many of

these adjacent communities require periodic fire to maintain 

their ecological structure and health (Kirkman and others 

1998b). Fires typically begin in pyrogenic longleaf pine 

forests and then spread into adjoining habitats such as seepage slopes, canebrakes, treeless savannas, and sand pine 

scrub. Without periodic fire, these communities, like longleaf pine ecosystems, change in ways that make them less 

suitable habitats for plants and animals that have evolved 

with fire. Continued degradation and eventual loss of longleaf pine forests from the southern landscape would not 

only be tragic for longleaf pine ecosystems, but could very 

well prove catastrophic for the numerous embedded biotic 

communities that are ecologically linked to them. The ecological imperative for restoring and sustaining these forest 

ecosystems is very clear.
9

logs using hand-powered pit-saws, which yielded only a few 

rough boards per day. By the 1700s, water-powered sawmills became common, but log transportation was inefficient 

and still largely confined to rivers (Frost 1993), with logging 

conducted on 5- to 6-km-wide strips along rivers, where logs 

could be dragged by oxen or horses and floated to the mill 

(Croker 1987). This so limited harvesting that, by the end of 

the Revolutionary War, most longleaf pine forests remained 

intact. After 1830, removal of longleaf pine accelerated 

significantly with the arrival of steam railroads, which were

soon followed by the use of steam skidders. By 1880, most 

of the longleaf pine along streams and railroads had been 

harvested (Frost 1993). During the next 40 years, the great 

forests of yellow pine were harvested, with temporary railroad spur lines laid down every quarter mile (Croker 1987). 

Skidders dragged logs to these spur lines, often destroying 

all trees too small to harvest, and left a scarred and mostly 

barren landscape. Longleaf pine forests were harvested from 

Virginia and the Carolinas south to Georgia and Florida, then

west through Alabama and Mississippi, into Louisiana and 

finally Texas. During 1896, 11.1 million m3 of yellow pine 

timber was cut and shipped to the northern United States and 

overseas markets (Mohr 1896). Timber extraction peaked 

in 1907, when 39 million m3 were removed (Wahlenberg 

1946). By 1930, nearly all old-growth longleaf pine was 

harvested, and lumber companies migrated west.

Extraction of naval stores by cutting wound faces in the bark

of longleaf pine trees began with the first European settlements (Frost 1993) and continued until the 20th century, when

it was supplanted by the petroleum industry. Because the 

pitch-soaked faces on these trees would readily ignite, many 

forests worked in this manner were destroyed by wildfire 

following abandonment. About two-thirds of the sites where

longleaf pine was harvested or burned by wildfires following 

naval stores extraction were later colonized by other tree 

species. Loblolly pine (a prolific seed producer) captured 

mesic Coastal Plain sites, slash pine invaded wetter flatwoods

areas, and shortleaf pine and hardwoods became dominant 

on upland sites. Irregular seed production, with good seed

years occurring at intervals > 5 years (Boyer 1990b), impaired

longleaf pine recovery and contributed to these losses. Even

where longleaf pine seedlings survived logging, they were

often consumed by feral hogs (Sus scrofa), causing many 

areas of potential longleaf pine forest to be lost (Schwarz 

1907). Large areas of longleaf pine forest were also converted to agriculture, beginning with early settlers who, 

like Native Americans, began by girdling trees and planting 

crops between dead standing snags. The settlers later burned

the snags and burned or dug out the stumps. Annual burning

to improve forage for livestock grazing frequently eliminated 

newly germinated longleaf pine seedlings. Between 1750 

and 1850, most of the more fertile longleaf pine sites were

converted to fields or pastures (Frost 1993), thereby removing

longleaf pine from the best upland areas. Although most of

the sandhills, flatwoods, and mountain soils are poorly suited

to agriculture, some were converted to pasture (Landers and 

others 1990). Florida was an exception, where sandhills sites 

are well suited for citrus fruit production (Mohr 1896). 

By the 1940s, harvesting second-growth longleaf pine became

an established practice, and construction of kraft-process 

pulp mills in the 1950s created demand for smaller trees. 

These developments accelerated conversion of naturally 

regenerated longleaf pine forests into plantations of species 

that grow more rapidly in the short term. Because of its 

slower early growth and lower survival rate, longleaf pine 

was seldom selected to reforest harvested lands. Thus, many 

second-growth longleaf pine stands on public and industrial 

lands were clearcut, mechanically site prepared, and planted 

with loblolly pine or slash pine. Nonindustrial private landowners often relied on natural regeneration following harvest,

and because insufficient longleaf pine seed was often present,

many of these sites regenerated into loblolly pine or slash 

pine forests. Old fields were also planted with these more 

rapidly growing species or colonized by them through seeding from adjacent areas. The result was a continuous decline

in the area occupied by longleaf pine ecosystems. Substantial future losses on private lands remain possible, because 

most of these longleaf pine stands consist of trees in the 

valuable sawtimber- and pole-size classes. 

Reduction in the frequency of fire further contributed to conversion of longleaf pine lands to other species. Extensive 

logging during the late 19th and early 20th centuries created 

very heavy loads of downed fuel, and this fuel supported 

numerous large wildfires that caused many areas to be devoid

of trees. Foresters then began to advocate excluding all fire

from the woods to protect young trees and allow for reestablishment of the forest. Although some individuals recognized

the natural and essential role of fire in longleaf pine ecosystems (Harper 1913), most people viewed fire as harmful. 

Since fire control practices aided in establishment of new 

forests, even though they were usually loblolly pine or slash

pine, these procedures were adopted as good forestry practices throughout the region (Frost 1993). 

Young hardwoods are quite susceptible to mortality from 

fire, and frequent fires typically limit hardwoods to a small 

stature in longleaf pine stands (Komarek 1977, Landers and

others 1990). Occasionally, random variation in fires or 

protective microsite conditions allowed hardwood stems to

survive several fires and become large enough to resist future

surface fires (Rebertus and others 1993). Thus, scattered 
10

hardwood trees occurred in the canopy or subcanopy of 

longleaf-dominated forests (Greenberg and Simons 1999). 

However, in the absence of fire, hardwoods are able to quickly

emerge from the understory and form a dense midstory that 

shades out herbaceous species and longleaf pine seedlings. 

Without frequent fire, hardwoods will ascend to eventually 

dominate the overstory, degrading sites not captured by 

other pines (fig. 3). Although the importance of fire in maintaining a healthy longleaf pine ecosystem is now widely 

recognized, many forests on private lands are still not burned 

regularly. Recent burning rates on private lands range from 

a high of 48 percent in Georgia to a low of only 15 percent 

in North Carolina (Outcalt 2000). Some stands are difficult 

to burn because they are close to urban areas and highways. 

Smoke from fires in such locations can have costly offsite 

effects. This problem is likely to grow worse as population 

growth creates more wildland-urban interface zones. Also, 

longleaf pine stands on private lands are small, and this 

makes burning them more expensive. Because of infrequent 

burning, many private lands containing longleaf pine are 

likely to suffer further habitat degradation. 

Since European settlement, longleaf pine lands have been 

lost to urban and residential uses. From 1987 to 1995, conversion of longleaf pine land in Florida to other uses resulted 

in loss of 37 000 ha of these ecosystems. During this 8-year 

period, ~ 3000 ha/year were converted from longleaf pine 

to urban uses, and 1500 ha/year were lost to agriculture 

(Outcalt 1998). Similar losses occurred in Georgia, while 

losses in North Carolina and South Carolina were at about 

half this rate. Future growth of the regional population and 

expected expansion of industrial plantations will likely 

result in a continuing decline of longleaf pine ecosystems on 

private lands. 

Social, Political, and Economic Context

Longleaf pine ecosystems have been very important in the 

Southeastern United States, providing an environmental 

setting and raw materials for social development in this 

region. Wild game, forage grasses, wood, and naval stores

were principal products of these forests (Franklin 1997). 

Figure 3—Former longleaf pine site invaded and occupied by oak.
11

During the early 20th century, affluent landowners, recognizing the value of longleaf pine forests as habitat for northern 

bobwhite and white-tailed deer, acquired large tracts to serve

as hunting plantations. Many large areas of longleaf pine exist

today because of the opportunities for hunting and timber 

harvesting provided by these lands. The appearance of these 

forests was also pleasing to many people, as it is today.

Although longleaf pine forests were valued by society, human

activities played a major role in their decline, with economic 

exploitation continuing until the future of these ecosystems 

appeared quite bleak. However, a combination of recent 

developments provides new hope that the negative trend for

longleaf pine forests can be reversed. Conversion of longleaf 

pine to other tree species has slowed as numerous Federal 

and State agencies have begun regenerating longleaf pine on

their lands following harvest. The presence of longleaf pine

on public lands has begun to increase as a result of concerted

efforts to establish new stands and restore degraded longleaf 

pine forests with fire and other appropriate techniques 

(Hilliard 1998, McMahon and others 1998). Interest in longleaf pine reforestation on private lands has surged recently 

because of incentives to private landowners provided by the 

Federal Government. From 1998 to 2000, longleaf pine was 

planted on 68 240 ha across the region. 

The southern forestry community has also gained an improved

understanding of longleaf pine ecosystems and come to 

appreciate the natural heritage that will be lost if restoration 

of these ecosystems is not undertaken. No single entity dominates land ownership in longleaf pine ecosystems, but a 

common sense of urgency among numerous groups has 

fostered partnerships that did not exist in the past. The Nature

Conservancy, Tall Timbers Research Station, Joseph W. 

Jones Ecological Research Center, U.S. Department of Agriculture (USDA) Forest Service, U.S. Department of Interior 

Fish and Wildlife Service, U.S. Department of Defense, 

Cooperative Extension Service, State agencies, private landowners, universities, and forest industry now work together 

to promote longleaf pine ecosystem restoration. In 1995, 

the Longleaf Alliance was formed to serve as a regional 

clearinghouse for a broad range of information about the 

regeneration, restoration, and management of longleaf pine

ecosystems. It facilitates communication among these groups

and provides training for private landowners who are interested in successful longleaf pine regeneration. 

The Federal Government uses financial incentives and technical assistance programs to promote restoration of longleaf 

pine ecosystems (table 2). Modest expansion of these forests

was realized when, for a 2-year period, longleaf pine was the

most frequently selected tree species for planting because of

incentives offered through the Conservation Reserve Program

(CRP). Federal tax laws also provide incentives for reforestation activities. In addition to these Federal programs, seven

Southeastern States have cost-share programs that encourage 

landowners to establish and maintain longleaf pine forests. 

All State forest management agencies in the region also offer

free management planning assistance to nonindustrial private

forest landowners. In aggregate, these public programs have

greatly encouraged establishment of new longleaf pine 

forests and contributed to the rehabilitation of degraded 

Table 2—Federal Government programs promoting restoration of longleaf pine ecosystems

Program Assistance, incentives, and purpose 

Conservation Reserve Program Financial incentives to landowners for planting and 

maintaining trees on designated nonforested land

Forestry Incentives Program Partial reimbursement for costs of establishing new forests, 

by means of natural or artificial regeneration, and costs of 

stand improvement

Forest Stewardship Program Education and technical assistance to landowners who 

manage forestlands for multiple resource benefits

Partners with Wildlife Program Cost-share assistance to landowners for restoring longleaf 

pine habitats

Safe Harbor Program Encourages voluntary habitat restoration by offering 

landowners incentives to preserve habitat of federally 

protected species 
12

longleaf pine ecosystems. However, recently adopted Federal

and State laws and regulations that are intended to improve 

or maintain air quality standards represent a serious challenge

to management efforts to burn these forests often enough 

to restore and sustain them. Mandated smoke management 

requirements restrict prescribed burning to a narrower set 

of conditions than in the past, thus reducing the number of 

possible burning opportunities during the year. 

Establishing longleaf pine is more expensive than establishing other southern pines. The price for bare-root seedlings 

of longleaf pine is about twice that of loblolly pine, because 

nurseries must grow longleaf pines at wider spacings to produce large seedlings suitable for planting. Seedling prices for

containerized longleaf pine are even higher, because greater 

labor and materials costs are involved. Planting costs are 

often somewhat higher for longleaf pine than for other pine 

species, because longleaf pine seedlings require greater care 

during the planting process. 

However, once established, longleaf pine becomes a lowrisk investment that is easily managed with prescribed fire, 

resistant to most pathogens and insects, and inexpensively 

renewed through natural regeneration. The superior growth 

form and wood quality of longleaf pine also produces excellent sawtimber. Typically 50 to 80 percent of the trees in 

forests of naturally regenerated longleaf pine are suitable for 

producing poles and pilings (Boyer and White 1990). For 

these reasons, longleaf pine can be marketed at stumpage 

prices 50 percent greater than those paid for the sawtimber 

of other southern pines. Longleaf pine forests can be managed to produce quality poles on a 40- to 60-year rotation 

(Franklin 1997). Also, recent evidence suggests that the rate 

of timber volume growth for longleaf pine is actually ~ 35 

percent greater than previously believed, which indicates 

that the past lack of appreciation for longleaf pine may have

been related to a systematic underestimation of its productive potential (Boyer 2001). In many areas, the high-quality 

pine straw (i.e., fallen pine needles) produced by longleaf 

pine provides additional income for landowners and processors, who market it as a landscape mulch (Williston and 

others 1990). 

Restoration Perspectives

Large-scale logging of native forests; conversion to agricultural, industrial, and residential uses; landscape fragmentation; invasion by aggressive species; and interruption of 

natural fire regimes have all contributed to the historical 

decline of longleaf pine ecosystems. This trend continued 

for many decades until the increasing threat to numerous 

rare plant and animal species and their habitats resulted in a 

growing social awareness that a very important part of the

South’s natural heritage was in jeopardy (Means and Grow 

1985, Noss and others 1995). Because longleaf pine still 

occurs over most of its natural range, albeit in isolated 

fragments, it is reasonable to conclude that restoration of 

these ecosystems is feasible (Landers and others 1995). 

Although the long-term negative trend has been recently 

slowed (reversed on public lands but continuing on private 

lands), resource assessments forecast a continuing overall 

decline without active intervention to reverse these losses. 

While the need for ecological restoration is clear, the choice 

of methods and the extent to which it should be pursued to 

ensure long-term ecosystem viability are less clear.

Defining Restoration

Although general definitions of restoration emphasize 

returning to a former state, it is not the goal of ecological 

restoration to recreate conditions that existed in North 

America prior to European contact in 1492. An essential 

element of all historically focused restoration approaches 

has been the description of a historically based reference 

condition, which enumerates the ecological attributes of the

native ecosystem (Covington and others 1997, Moore and

others 1999, Palik and others 2000, Wagner and others 2000).

The degree to which restoration objectives are achieved 

may then be assessed by mathematically comparing existing 

community composition with that in the reference condition 

(Provencher and others 2001). Although such approaches 

are well intended, returning an ecosystem to pre-European 

conditions or any other arbitrary point in history is both 

impractical and impossible for many reasons. Since the Age 

of Exploration and the Industrial Revolution, human impacts 

on North American ecosystems have led to extensive landscape domestication and fragmentation. Human activity is 

also responsible for the widespread occurrence of exotic 

species and extirpation of many native species. Early records 

do not provide reliable, detailed data about the composition, 

structure, and processes in indigenous ecosystems (White 

and Walker 1997). Restoration efforts based on such limited 

information would overlook key ecosystem components and 

processes. 

Ecosystem variability is so strongly dependent on scale, or

affected by humans, that identification and application of 

meaningful and relevant historical data for management is

essentially impossible. Poorly conceived reference conditions could be used by naive scientists and managers to define

a misguided, impractical, or expensive template for the future

(Parsons and others 1999). Much of what European immigrants perceived as “pristine wilderness” was actually occupied for > 12,000 years by millions of native people who, 

although living in preindustrial societies, actively managed 
13

their landscape through extensive use of fire and other available technology (Pyne 1997, Pyne and others 1996). By the

time significant numbers of immigrants arrived, many of 

these lands were depopulated of their native inhabitants by

newly introduced diseases and occupied by woody vegetation that had invaded former village sites and abandoned 

agricultural fields during the ensuing decades (Stanturf and 

others 2002). Thus, much of what is recorded in the journals 

of early explorers, naturalists, and settlers is the product of 

both natural processes and the actions of Native Americans 

(Carroll and others 2002). 

Finally, the ecosystems of today are substantially different 

from those of the past not only because of biological alteration, but also because their physical environment has slowly

changed. Natural variations in climate have influenced the 

establishment and development of forests for millennia. 

Many forests were initiated during periods when temperatures were much cooler. The composition, structure, and 

process dynamics of forests that thrived during cooler 

climatic periods such as the Little Ice Age (1350 to 1850) 

probably differ from those forests that developed during 

the 20th century, when temperatures became significantly 

warmer (Millar and Woolfenden 1999). Therefore, while 

it may not be desirable or even possible to restore forest 

ecosystems with historical authenticity, it is quite possible 

to restore them with natural authenticity; that is, to restore 

them to a condition in which compositional, structural, and 

functional components are present within an appropriate 

physical environment that sustains the ecological processes 

essential for (1) native species perpetuation and evolution, 

(2) ecosystem resiliency to disturbance and adaptation to 

long-term environmental change, (3) providing goods and 

services for human societies, and (4) harboring rare and 

endangered species and otherwise conserving biological 

diversity (Clewell 2000).

Perhaps the broadest definition of ecological restoration 

is “an intentional activity that initiates or accelerates the 

recovery of an ecosystem with respect to its health, integrity 

and sustainability” (Society for Ecological Restoration 

International 2002). The National Research Council explains 

ecological restoration in more detail as “the return of an 

ecosystem to a close approximation of its condition prior 

to disturbance; both the structure and the functions of the 

ecosystem are recreated; merely recreating the form without 

the functions, or the functions in an artificial configuration 

bearing little resemblance to a natural resource, does not 

constitute restoration; the goal is to emulate a natural, functioning, self-regulating system that is integrated with the 

ecological landscape in which it occurs” (Cairns 1995). These

and other definitions raise a number of questions, not the 

least of which are, “What were predisturbance forest and 

landscape conditions like, and how have they changed?” and 

“In view of these changes, how can restoration actions most 

effectively establish a biotic community that is now suited 

to the prevailing environment?” Although restoration can 

never legitimately be a return to the past, an awareness of 

historical alteration of biota, previous erosion of soils, and 

former fluctuations and current trends in climate are among 

the many factors that must be considered. Much like trying 

to hit a moving target, attempting to restore a previously 

existing historical ecosystem is less important than matching 

plant species to site conditions, reestablishing essential 

natural processes, and mitigating potential threats to the 

current and future ecosystem (Allen and others 2001). 

Measuring Success

Measuring success, with restoration approaches that utilize a

vaguely defined historical reference condition as the target,

creates inherent difficulties for practitioners. In most instances,

appropriate reference areas are largely unavailable, and those

that do exist are often degraded by impaired ecological processes and/or past human disturbances that are not readily 

apparent. Thus in selecting a site as a historical reference 

area, the individual may, in fact, be choosing a site that is 

partly or even wholly an artifact of human disturbance on a 

temporal and/or spatial scale that is not easily recognized. 

Monitoring schemes which use a historical reference condition to evaluate the progress of restoration typically focus 

on species similarity, with metrics that merely evaluate the 

degree to which the biotic community on the reference site 

is replicated on the site being restored. Little if any regard is

accorded the unique ecological attributes of each site that

influence community development. Monitoring and evaluation of this type represents a nondynamic approach to restoration that is reflective of a single-condition ecosystem model

and analogous to outmoded notions like “climax” and “the

balance of nature.” Such a static approach will not adequately

account for natural disturbance, disequilibrium, and dynamic

change in the ecosystem. 

Restoration approaches that emphasize the functional nature 

of ecosystems will likely be more feasible alternatives by 

reestablishing biota that are compatible with the dynamic 

changes in present and future ecosystems. Key attributes 

to be restored include (1) species composition, (2) vertical 

structure, (3) horizontal pattern, (4) spatial heterogeneity,

(5) properly functioning ecological processes, and (6) ecosystem resiliency sufficient to permit recovery from disturbances

at multiple scales (Hobbs and Norton 1996). Although the 

restoration process must identify and mitigate to the degree 

possible forces that threaten ecosystem integrity, looking 

beyond historical conditions, it must also accommodate those
14

factors of change that cannot be mitigated and establish a

community that is suitably adapted to prevailing environmental conditions. Monitoring is essential and must include data

gathering, analysis, and meaningful interpretation that produce useful feedback for the adaptive management process. 

Although monitoring need not always be sophisticated to be 

effective, numerous variables related to (1) plant and animal 

communities, species, and populations; (2) soils, hydrology, 

and water quality; (3) landscape patterns and functions; and 

(4) ecological processes and response to disturbance events 

can be measured (Allen and others 2001). The values of 

such variables may then be compared with values falling 

either inside and outside the range of natural variability 

(Hobbs and Norton 1996). 

In an integrated analysis of structure and function, monitoring data are compared with values for ecosystem attributes 

that facilitate the functioning of natural processes, including 

recovery from disturbance. Recognizing the widely variable

nature of natural longleaf pine ecosystems, the acceptable 

range of values for many of these attributes will be quite 

broad. Structurally, success should strive for an overstory 

dominated by longleaf pine, occurring as uneven-aged stands

or even-aged patches across an uneven-aged landscape 

mosaic. Depending on site type and location in the native 

range, a smaller component of other tree species, such as

slash pine or oaks, may be present. These may occur singly

or in clusters. Generally, the midstory should be absent or

mostly composed of ascending longleaf pines. The understory should be dominated by native grasses and forbs, and

few shrubs and vines should be present. Because the composition and abundance of graminoids varies greatly throughout

the native range of longleaf pine, specific quotas for representation of individual species (e.g., a specific percent cover

for wiregrass) are inappropriate. Functionally, success should

consider ecological processes such as periodic surface fires,

natural regeneration that leads to normal stand replacement 

dynamics, nutrient cycling that maintains primary productivity, and suitable habitat that facilitates life cycle completion by numerous native organisms. Reductions in habitat 

fragmentation, population isolation, and species rarity will 

be achieved by augmenting existing longleaf pine fragments 

and creating new habitat patches, increasing connectivity 

among existing fragments and new habitat patches, and 

facilitating the role of fire at local and landscape scales to 

improve habitat quality and enhance biological diversity.

Future Ecosystems

Sole reliance on natural processes may require decades or

even centuries to restore degraded ecosystems. Where threatened ecosystems need urgent attention, restorationists must

identify environmental problems and intervene with remedial

treatments (Dobson and others 1997). Although factors that

are unknown or beyond their control may frequently cause

restorationists to accept partial solutions to the challenges 

confronting them, inability to completely restore ecosystems

does not diminish the importance of restoring the understood

ecological functions and resource values. Using an holistic 

approach, ecological communities should be established, to

the greatest degree possible, that not only resemble original

forests but are also well suited to prevailing and future environmental conditions (Allen and others 2001). In longleaf 

pine forests, the restoration focus is on reestablishing the

natural ecosystem structure and function through (1) adjusting composition by favoring native species and suppressing 

undesirable species, (2) modifying vertical and horizontal

structure at the local and landscape scales to provide a full

range of habitat conditions that are appropriately interconnected, and (3) ensuring that ecological processes such as

periodic fire and natural longleaf pine regeneration remain 

fully operational at multiple spatial and temporal scales. 

Restoration is a long-term incremental process of serial 

approximation, where all gains are valued as forests are progressively transformed from degraded to healthy conditions. 

Successful restoration requires more than just establishing 

overstory trees and must also include improving, and establishing where necessary, understory plants and all other 

appropriate ecosystem components. It may often be best to 

restore a vigorous cover of native understory plants before 

establishing the overstory. 

Encouraged by ecosystem management policies, cooperative 

partnerships between research scientists and land managers 

have facilitated a more effective application of ecological 

knowledge and silvicultural techniques for addressing forest

ecosystem restoration challenges in an adaptive management

framework. As application of ecological theory has benefited

land management, the practice of ecological restoration will 

continue to provide insights into the way that ecological 

communities are assembled and ecosystems function (Dobson

and others 1997). A relatively recent advance in this realm 

has been a greater appreciation for the importance of ecological thresholds and an improved understanding of nonlinear

succession processes that appear to be highly relevant to disturbance-prone ecosystems (Friedel 1988, 1991; Westoby 

and others 1989). Since transforming a degraded ecosystem

into a more desirable condition through accelerating biological change or reinitiating succession is the principal paradigm of restoration ecology, the restoration process requires 

redirecting development of the ecosystem along a trajectory 

that is assumed to have existed prior to disturbance (Aronson

and others 1993). However, ecosystems under stress do not 

always exhibit an orderly gradual development as suggested 
15

by classical succession theory (Clements 1916), but often 

undergo rapid transitions among multiple metastable states 

(Drake 1990, Hobbs 1994). These transitions are indicative 

of a nonlinear successional process with threshold responses 

to management activities and environmental factors, especially those that generate acute or chronic disturbances. 

The occurrence and persistence of particular ecosystem states

depend on specific combinations of driving forces, and once

an ecosystem proceeds beyond a threshold condition, massive

management inputs may be required to restore it to its former

state (Hobbs and Norton 1996). Restoration may be viewed 

as attempting to force transitions toward a more desirable 

state, an effort that requires knowledge about the factors that 

must be manipulated to successfully cross or recross these 

thresholds. The fact that alternative stable states are possible 

for any individual location even under natural conditions 

should cause us to reflect carefully when establishing goals 

for restoration and sharply calls into question restoration 

efforts based on nondynamic, historically referenced, singlecondition ecosystem models (Brockway and others 2002). 

When a degraded ecosystem has not yet crossed a critical 

threshold, removal of the degrading influence may be all that

is required to encourage restoration; however, if an important

threshold has been crossed, then simply removing the degrading influence will not be sufficient to allow transition back 

to an approximation of the former state (Hobbs and Norton 

1996). Rather, active intervention will be required. Critical 

thresholds in terrestrial ecosystems have been linked to the

magnitude, frequency, and sequencing of multiple disturbances in the physical environment (i.e., windstorms, floods, 

fire, drought, soil erosion). These thresholds may also involve

important changes in functional groups of plants, with transitions between states being much more difficult when one 

functional group is largely displaced by another (Hobbs 

1994, 1996). 

A fundamental challenge for restoration ecology is the problem of developing practical and cost-effective methods for

forcing these transitions (Hobbs and Norton 1996). An ecological model, using principles from the theory of multiple 

stable states, has been proposed for restoring longleaf pine

ecosystems, without returning them to presettlement conditions (Walker and Boyer 1993). Such an approach could go 

a long way toward reestablishing continuous forests similar 

to the Dauerwald of Europe, in which naturally regenerating 

indigenous forest ecosystems have been restored on suitable 

sites and are sustained along with other native species in an 

uneven-aged landscape matrix that provides high-quality 

habitat for conserving biological diversity and goods and 

services for human societies (Schabel and Palmer 1999). 

Restoration Methods

Restoration Framework

Recent research has produced a wealth of knowledge and

experience about the restoration of longleaf pine ecosystems.

Valuable restoration techniques are also being developed by

forestland managers who are making observations and using

monitoring procedures in successful application of adaptive

management strategies. Despite these advances, the remaining

uncertainty has fostered debate about the best approaches for

restoring longleaf pine ecosystems. This debate has focused 

on whether restoration is best achieved by (1) a passive 

approach, (2) restoring structure, (3) restoring function, or

(4) restoring both structure and function. The passive or nonmanipulative approach is based on the belief that protection 

from all natural and human disturbances is the best alternative and was the chosen option for much of the 20th century, 

when fire exclusion was a major emphasis. The restoring 

function approach is tested by use of prescribed fire in longleaf pine forests and is based on the idea that fire is the principal natural driving force for development and maintenance 

of these ecosystems. The restoring structure approach is 

tested by using mechanical treatments to remove invasive 

woody plants and is based on the idea that an ecosystem that 

is outside its range of natural variation requires intervention 

to restore forest structure and ecosystem health. The structure and function approach employs both mechanical treatments to manipulate structure and reintroduction of fire as 

an ecological process. 

Desirable changes in longleaf pine communities can be 

achieved by using a variety of methods, machines, and 

products, singly or in combination. Physical or mechanical 

treatments include complete overstory harvesting, selective 

thinning of overstory and midstory trees, and shredding or 

mowing midstory and understory plant layers. Chemical 

treatments, principally herbicide application, can be used to 

selectively reduce undesirable plants. Prescribed fire may

also be used to reduce midstory, understory, and occasionally overstory layers and encourage fire-tolerant plants. In

fact, frequent fire is crucial for ecosystem restoration, and the

use of other treatments should be planned so as to facilitate 

the eventual application of prescribed fire. In highly degraded

ecosystems, biological approaches such as reintroducing 

extirpated species will likely be required for full restoration. 

Each type of treatment could be appropriate for restoring 

particular longleaf pine ecosystems, depending on their 

current condition, location, and ownership. 

Selecting Techniques

Historical events and changing patterns of land use have 

resulted in an array of sites in various conditions that are 
16

candidates for restoration of longleaf pine ecosystems. While

about 1.2 million ha currently support an overstory of longleaf pine (Outcalt and Sheffield 1996), only 0.5 to 0.8 million

ha of these lands have intact native understories (Noss 1989).

Additional candidate areas exist that have overstories with

little longleaf pine and understories that range from having

most of the native species to some that are highly altered by

past disturbance (Outcalt 2000). This variety of conditions 

exists throughout the range of longleaf pine, on sites from

dry sandhills to wet savannas. Suitable restoration techniques

depend on the site type and degree of ecosystem degradation 

(table 3). The types of longleaf pine ecosystems discussed 

in the following section on restoration prescriptions are 

based on the classification of Peet and Allard (1993), with 

sandhills corresponding to their xeric and subxeric series, 

flatwoods and wet lowlands to their seasonally wet series, 

and uplands to their mesic series. However, we include their 

Piedmont/upland subxeric woodland community in the 

uplands rather than sandhills. 

Restoration Prescriptions

Exclusion of fire from many existing longleaf pine forests 

allowed forest fuels to accumulate to hazardous levels and 

permitted substantial expansion of understory shrubs and 

hardwoods. Although the introduction of dormant-season 

prescribed burning reduced the fuel buildup, midstories were 

often so far advanced that they were only minimally affected 

by these cool-season fires. To restore these ecosystems more

fully, it is necessary to reintroduce growing-season fire, the

ecological process that most effectively creates and maintains the structure, composition, and function of such forests.

Growing-season burning is usually conducted from March 

to July, with the period from late August through September 

avoided to prevent excessive mortality among young longleaf pines (Robbins and Myers 1992). 

Some longleaf pine forests, such as those on wet sites (i.e., 

flatwoods), may require biennial burning over a period of

several decades to substantially improve understory composition and structure. Where it took > 30 years of dormantseason burning (every 4 to 6 years) for the understory to 

reach its current status, there is no reason to expect that this

condition can be reversed with a single fire or during a 5-year

period. As long as the sequence of treatments is progressively

changing the community along a desired trajectory, then 

progress is being made toward eventual restoration.

Xeric and subxeric sandhills dominated by longleaf pine 

with native understory—In many existing longleaf pine

forests, the lack of frequent fire allowed turkey oak (Q. laevis),

bluejack oak (Q. incana), sand live oak (Q. geminata), and

sand post oak (Q. stellata var. margaretta) to develop into a

scrub oak midstory. However, because moisture and nutrients

are very limited on these sites, they do not develop a continuous closed canopy of midstory hardwoods even in the 

absence of frequent fire. Therefore, some of the understory 

grasses survive. These grasses and the needle litter from longleaf pines can carry at least a patchy prescribed fire. Repeated

Table 3—Prescriptions for restoring longleaf pine ecosystems in varying degrees of degradation

Moderately degraded Very degraded Highly degraded

Overstory: Longleaf pine Other trees Other trees

Understory: Native plants Native plants Nonnative plants

Xeric and subxeric sandhills: Growing-season fire Mechanical harvesting Roller-chop twice and burn

Dormant-season fire Growing-season fire Herbicide if needed

Mechanical removal and Herbicide sprouts Plant LLP seedlings

 herbicide hardwoods Plant LLP seedlings Sow native understory seed

Montane and mesic uplands: Growing-season fire Growing-season fire Growing-season fire

Dormant-season fire Mechanical harvesting to Harvest, chop, harrow

Mechanical removal and create canopy gaps Herbicide if needed

 herbicide midstory Plant LLP seedlings Plant LLP seedlings

Sow native understory seed

Flatwoods and wet lowlands: Growing-season fire Growing-season fire Roller-chop twice and burn

Dormant season fire at Mechanical harvesting Herbicide if needed

 2-year intervals Roller-chop once and burn Plant LLP seedlings

Plant LLP seedlings Sow native understory used

LLP = longleaf pine.
17

applications of fire during the growing season are effective 

in restoring these sites by gradually reducing the density 

of the midstory scrub oaks (Glitzenstein and others 1995). 

Fire causes wounds on the stems of these hardwoods, and 

these wounds are enlarged by subsequent fires until the stem 

is girdled, the top dies, and the trunk breaks. Sprouts arise 

from some top-killed stems, but their growth is curtailed by

recurrent fire. This process is aided by increases in understory

grasses that are stimulated by fire to produce more biomass 

that becomes fuel for future fires. Fires stimulate grasses and

forbs to produce flowers and seeds (Christensen 1981, Clewell

1989, Outcalt 1994, Platt and others 1988a), that aid in colonization of newly exposed microsites. Although some of the 

hardwood stems survive, this is not undesirable since they 

are a natural part of longleaf pine ecosystems (Greenberg 

and Simons 1999). Restoration aims to decrease the number 

of midstory hardwoods, not completely eliminate them.

Reintroducing growing-season fires into xeric longleaf pine 

forests that have not been burned for a prolonged period may

result in increased mortality among older trees during the 1-

to 3-year interval following burning. The cause of this is not 

known, but it seems to be related to excessive accumulation 

of forest litter around the base of larger longleaf pines and

damage to roots or cambium or both as a result of smoldering combustion of this litter. To decrease this mortality, a

series of dormant-season fires at the shortest intervals fuel

levels should be applied to reduce the litter buildup. Growingseason fire may then follow, when fuel is sufficient. In all

prescribed burning, duff moisture levels must be high enough

to prevent ignition of the litter layer at the base of larger 

longleaf pines, in order to avoid excessive fine-root damage 

and girdling of trees by cambial injury at the root collar. In 

many areas, the duff layer around the base of large trees is 

dry down to the mineral soil in the spring, during the early

part of the growing season. Summer rains then occur, the

drought index drops, and conditions seem suitable for prescribed burning. However, if rainfall has not been sufficient 

to completely re-wet the duff layer at the tree base, enough 

heat will be generated by a prescribed fire to evaporate the 

modest amount of moisture that is stored in the upper litter 

layer. The litter will then ignite and burn slowly for many 

hours, producing temperatures sufficient to cause cell death 

in roots and the base of the tree. Thus, it is important to 

determine whether the lower duff layer (i.e., the part that 

is in contact with mineral soil) at the base of larger trees is 

sufficiently wet. If the lower duff is wet and the upper duff 

is dry, fire should consume the dry top layer while the wet 

lower layer protects the roots and root collar. This favorable 

wet duff condition may not occur until late in the growing 

season during some years, because it takes significant precipitation to re-wet this layer once it has been dried completely. 

If the duff layer is thick and precipitation is below normal, 

this favorable condition may not occur until the dormant 

season in some years. Thus, it seems prudent to use a series 

of dormant-season fires to gradually reduce the accumulated 

litter before switching to growing-season burning. However, 

some older longleaf pine will die even if prescribed burning 

is done carefully. Studies underway at Eglin Air Force Base 

in Florida are aimed at identifying the factors most critical 

in predicting tree mortality and should provide information 

useful in refining burning prescriptions to minimize such 

losses.1 Managers should be prepared for some mortality 

among older longleaf pine trees, because completely eliminating such mortality in long-unburned ecosystems may not 

be possible. Fire-killed trees may be salvaged to obtain some 

economic benefit or retained onsite as snags to enhance wildlife habitat. The ecological negative effects of not burning 

longleaf pine forests will far outweigh the loss of some 

older trees. Delays in reintroducing prescribed burning only 

increase the likelihood that these ecosystems will be lost as 

a result of catastrophic wildfire and invasion by aggressive 

pines and hardwoods. 

Although sandhills where scrub oak expansion is the major 

problem tend to respond quickly after three or four growingseason fires, application of supplemental treatments can 

accelerate the restoration process. Mechanical methods such 

as chain saw felling, girdling, or chipping on site can reduce 

midstory hardwoods (Provencher and others 2001), and following these with prescribed fire will stimulate grasses and 

forbs and reduce the growth of hardwood sprouts. If woody 

material from the midstory is not chipped or removed from 

the site, it must be allowed to decay before prescribed fire 

is introduced, so that it does not add to fuel levels. These 

mechanical treatments can be expensive and are most appropriate for critical areas in need of rapid restoration, such as 

red-cockaded woodpecker colony sites or areas along the 

wildland-urban interface where it is difficult to schedule the 

series of prescribed fires required for restoration. 

Hexazinone herbicide can also be useful in accelerating the 

restoration process, although it may significantly contribute 

to onsite fuel levels if used in areas containing high numbers 

of large target species. It is very effective in decreasing midstory hardwoods, with little or no short-term reductions in 

understory grasses and forbs on sandhills sites (Brockway 

and others 1998). The rate of restoration is significantly more

rapid when hexazinone treatment is combined with prescribed

burning than when only prescribed burning is employed. 

The combined treatment can produce the desired results with

1 Personal communication. 2002. J. Morgan Varner, Graduate Student, 

University of Florida, 118 Newins-Ziegler Hall, Gainesville, FL 32611.
18

the first fire following hexazinone application (Brockway 

and Outcalt 2000). Because hexazinone does affect woody 

species, the cover of desirable nontarget species, gopher 

apple (Licania michauxii), for example, may be reduced 

for a time. However, hexazinone application is a one-time 

treatment intended to restore an area that can subsequently 

be maintained easily by periodic prescribed fires that mimic 

the natural fire regime. Application rates of 1 to 2 kg a.i./ha 

will produce 80 to 90 percent oak mortality without longterm damage to herbaceous understory species. When the 

liquid formulation is applied in a 2- by 2-m grid pattern, only

a very small portion of the soil surface is treated directly. 

This minimizes damage to grasses and forbs, but kills many

oaks. During dry periods, however, hexazinone may photodegrade before there is enough rainfall to carry it into the

soil where the oak roots absorb it. This problem was encountered by managers at Eglin Air Force Base, where very little

or spotty oak mortality was observed after application 

(Berish 1996). This problem can be avoided by applying 

hexazinone during periods prior to anticipated precipitation. 

Granular hexazinone is less subject to this problem because 

it will lie on the soil surface, without significant loss in 

strength, until rain arrives. However, granular hexazinone 

does have the potential to cause a greater reduction in the 

cover of grasses and forbs when it is applied uniformly over

the entire site (Brockway and others 1998). Broadcast application must also be conducted with care to avoid distribution 

overlap that could double the applied rate and create strips

where no understory plants survive. Oak roots are extensively distributed in the soil, and it is better to miss a small 

area between strips than to overlap strips during application. 

If some larger oaks are to be retained, hexazinone application

should be kept > 30 m away from these trees, because of 

their widely spreading root systems. Even at this distance, 

some oak trees may be killed. Leaving completely untreated 

areas of 0.5 to 1 ha may be the most effective way of creating

refuges for larger oaks. 

Xeric and subxeric sandhills dominated by other trees 

with native understory—Large areas exist where scrub oaks

have become dominant following the harvesting of longleaf 

pine. Although somewhat suppressed in the absence of frequent fire, the understory plant community in such areas still

contains many of the native species. There are also many 

areas that were converted to slash pine plantations following 

the removal of longleaf pine. Although understory species, 

especially the important grasses, can be killed when the soil

is disturbed on dry sandhills sites (Grelen 1962, Outcalt 1983,

Outcalt and Lewis 1990), some of these areas also have fairly

intact understory communities. These understories survived 

largely because site preparation was not intensive or because

of fortuitous rainfall and higher soil moisture levels following

soil disturbance. There are also extensive areas in western

Florida where Choctawhatchee sand pine (P. clausa var. 

immuginata) invaded former longleaf pine lands following 

harvest. Unlike slash pine, sand pine is well adapted to dry

sites, where it forms nearly continuous canopies that severely

reduce understory density. However, plant diversity in these 

stands is generally unaffected, with native species surviving 

but being less numerous (Provencher and others 2001). 

Restoration under these conditions requires invigorating the 

herbaceous understory, reducing the scrub oak tree layer, 

removing slash pine or sand pine if present, and establishing 

longleaf pine seedlings. 

Areas dominated by scrub oak are often difficult to burn 

because fuel is sparse and not contiguous. These sites can be

improved by mechanical treatment with a small (3 to 5 t) 

single-drum roller-chopper with no offset. Heavier choppers 

with offset rollers should be avoided because they can cause

excessive soil disturbance that will harm understory plants.

The objective of this treatment is to knock down the oaks and

compress them into a ground layer that, after drying, will 

carry a prescribed fire. By contrast, slash pine plantations 

often have enough needle litter to support a prescribed fire.

Burning these plantations will invigorate the grasses, allowing them to accumulate root reserves and thereby increase 

their ability to recover from the adverse effects associated 

with removal of the slash pine and establishment of longleaf 

pine seedlings. A second fire following harvest will remove 

logging slash, help control oak sprouts, and increase the 

cover of herbaceous species. If scrub oaks are a serious problem, hexazinone can be applied as outlined earlier. Application can be made prior to harvest, in which case the logging 

activity will knock down many of the standing dead stems, 

which will then serve as additional fuel for prescribed fire. If 

herbicide is applied after logging, it is best to let most dead 

oak stems fall over before burning, as this will help remove 

debris prior to planting of longleaf pine seedlings. Sand pine 

often grows so densely that it must be removed to release 

surviving understory species. Sites can then be burned to 

remove logging slash, reduce abundant sand pine seedlings, 

and consume sand pine seed. 

Options for establishing longleaf pine seedlings include 

planting of bare-root and containerized seedlings by manual 

or machine methods (Barnett 1992, Barnett and McGilvray 

1997, Barnett and others 1990). Site preparation, other than

that discussed above, should be avoided to protect the understory plant community. It is much less expensive to plant 

additional longleaf pine seedlings to compensate for lower 

survival than it is to reestablish key understory species lost 

as a result of excessive soil disturbance. If grass competition 

is vigorous (> 60 percent cover) and bare-root seedlings are
19

being used, a planting machine with a small scalper blade 

can be used to increase seedling survival (Outcalt 1995). 

Although this removes a strip of vegetation ~ 1 m wide, native

grasses and forbs will recolonize these strips within 3 to 5 

years, if invasive woody plants are discouraged by periodic 

growing-season fire. Planting containerized longleaf pine 

seedlings on sites prepared only by burning results in acceptable survival rates, but hexazinone application may increase

survival on areas with vigorous scrub oak competition. 

Xeric and subxeric sandhills without native understory—

These sites once supported native longleaf pine ecosystems, 

but severe disturbance has altered their vegetation greatly. 

They either have no longleaf pine trees and a much altered 

understory, or longleaf pines have been reestablished but the

native understory has not. Most of these sites were once used

for agriculture or intensively managed forest plantations. 

Restoration of the once diverse understory is a formidable 

and expensive task on such sites. We do not know a great 

deal about restoring understory plant communities; in this 

area our experience is limited to a very few operational-scale 

restoration projects. In most cases, the first step is removal 

of trees other than longleaf pine from the overstory. Since 

there are few understory plants to protect, many options are 

available for site preparation. Chopping with a double-drum 

offset roller-chopper effectively controls all competition and

produces a clean site for restoration (Burns and Hebb 1972). 

This treatment can be combined with burning if there are 

significant quantities of woody residue. Because much of 

the nutrient capital on these sites is in the litter layer and 

upper soil horizon, any root raking and shearing must be 

done carefully so that soil and litter movement is minimized. 

Longleaf pine bare-root or containerized seedlings can be 

planted on the site after the soil has settled. 

Understory plants should be restored while longleaf pine 

seedlings are being planted, because the site treatments associated with planting reduce competition and increase onsite 

operability. The most critical part in this process is the reestablishment of grasses, because of their importance as fuel to

support recurrent fire. To date, most work has been in the

eastern portion of the range and has focused on reestablishment of wiregrass (Means 1997, Seamon 1998). A native

bunchgrass, wiregrass has an average density of 5 bunches/m2

in healthy longleaf pine ecosystems (Clewell 1989). Wiregrass 

seed has been collected efficiently, and its seedlings have

been grown and planted successfully using recently developed technologies. Planting wiregrass seedlings at the high

densities typical of natural stands is too expensive and fortunately unnecessary. Once established, wiregrass bunches

will expand vegetatively and, if frequently burned, will produce seed that can aid in spreading the grass to unoccupied 

areas. Thus, a planting density of 0.5 to 1 seedling/m2

 can be

used to successfully reestablish this species (Outcalt and

others 1999). Wiregrass plugs can also be planted successfully under existing plantations of longleaf pine (Mulligan 

and others 2002). 

Any existing hardwood midstory must first be removed by

repeated burning, mechanical felling, or herbicide application. A heavy-duty woods-harrow is then used to disk strips

between trees. In the spring, wiregrass plugs can be planted 

at a 1- by 1-m spacing in these strips. Application of fertilizer during the second or third growing season will stimulate wiregrass growth (Outcalt and others 1999). Fertilizer 

should be applied only around wiregrass plants to avoid 

stimulating the growth of competing vegetation. In pastures 

occupied by bahia grass (Paspalum notatum), cultivation 

will break up the old sod, and use of herbicide will improve 

both the survival and growth of wiregrass (Uridel 1994). 

Direct seeding can also be used to reestablish wiregrass 

between rows of trees in newly planted and existing plantations. This method is less expensive than growing seedlings 

or planting plugs (Hattenbach and others 1998). Small quantities of seed can be collected by hand or with a hand-held

seed stripper. A tractor-mounted flail-vac is useful for collecting larger quantities of seed. Seed can also be collected by

harvesting the entire seed stalk. If entire seed stalks are 

collected, they should be spread on the restoration site soon

after harvest to prevent seed loss from heating and fungal 

growth. Seed can be spread manually or with a small bale

chopper. Seed or seed stalks can be collected from native 

understories that are purposely burned during the growing 

season to stimulate flower and seed production. The collection of seed from native understories diversifies the seed 

mix because it includes the seed of other native species that 

have mature seed at the time of collection. Reestablishment 

through seeding is more versatile than planting, because 

seeds can either be stored in woven bags or taken directly 

to the site and sown. Rolling sown seed into the soil can 

improve wiregrass establishment and survival (Hattenbach 

and others 1998). Other grass species are also part of the 

native understory in sandhills longleaf pine forests and 

should be included in seed mixes. Pineywoods dropseed 

(Sporobolus junceus), for example, is quite common on many

sites. It, like wiregrass, will produce seed following fire. 

Its seed can be collected by hand and mixed with wiregrass 

seed for sowing on restoration sites. Preliminary trials 

suggest that more pineywoods dropseed seedlings than wiregrass seedlings become established if equal amounts of seed 

of these species are sown. In addition, many of the young 

dropseed plants produce seed during the first year following 

establishment. 
20

Longleaf pine forest understories are of course more than

just grass. Smith and others (2002) compared the understories of remnant xeric longleaf pine areas with those of 

plantations on old-field sites established 30 to 40 years ago. 

Although the remnant longleaf pine sites did have a higher 

species diversity, nearly 90 percent of the understory species 

in the plantations were native to natural longleaf pine communities. Similar comparisons for the sandhills of South 

Carolina showed that species abundance was the same in 

plantations and reference stands, except that wiregrass and 

dwarf huckleberry (Gaylussacia dumosa) were significantly 

reduced in plantations (Walker 1998). These and other 

restoration studies (Hattenbach and others 1998) strongly 

suggest that many of the understory species on sandhills 

sites either survive extreme disturbance as propagules in the 

soil or are able to reinvade sites after the disturbance ends. 

Thus, restoration does not require that every plant species 

be reintroduced. Nevertheless, it will probably be necessary 

to reintroduce certain common species that do not easily 

reinvade or survive and some rare species (Glitzenstein and 

others 1998, 2001; Walker 1998).

Flatwoods and wet lowlands dominated by longleaf pine

with native understory—As with other site types, restoration on these lower, wetter areas is designed to restore 

diversity to longleaf pine communities that have an understory that has been captured by woody species and has in 

many cases developed a substantial midstory layer. The 

management objective is to reduce woody understory and 

midstory species and allow grasses and forbs to increase and

eventually become dominant. Prescribed fire is one tool for

accomplishing this transition, with growing-season fires

being at least as useful as and often more effective than

dormant-season burns for readjusting understory composition. Although growing-season fire should be favored, 

dormant-season fire may be necessary to reduce fuel loads 

before initiating growing-season burning. One or two 

dormant-season fires may be used to gradually reduce litter 

buildup before the first growing-season burn is applied. 

These dormant-season fires should occur within a 2-year 

period to minimize accumulation of fuel between burns. 

Miller and Bossuot (2000) recommend that initial burns be 

conducted when the Keetch-Byram Drought Index (KBDI) 

is < 250 (Keetch and Byram 1968). Flatwoods understories 

dominated by saw palmetto (Serenoa repens), gallberry 

(I. glabra), wax myrtle (Myrica cerifera), and sweetgum 

(Liquidambar styraciflua) are quite resistant to fire. Only 

repeated fires at short return intervals over a long period 

significantly reduce these woody species (Waldrop and 

others 1987). Thus, burning every 2 years for a period of 

10 to 20 years may be required to readjust the understory 

composition on wet sites. 

Managers at Myakka River State Park in Florida have had 

some success using lightweight choppers or heavy-duty 

mowers to reduce saw palmetto coverage and dominance 

(Huffman and Dye 1994). Both of these mechanical methods

cause very limited soil disturbance and thus do not reduce 

native grass species. Preliminary findings from research 

underway indicate that the chopping treatment is more effective for reducing saw palmetto cover. Prescribed burning 

3 to 6 months before or after these mechanical treatments 

seems to increase their effectiveness.

Flatwoods and wet lowlands dominated by other trees 

with native understory—These sites are a mixture of 

naturally regenerated stands that were invaded by slash pine 

and loblolly pine after the removal of native longleaf pine 

and plantations that were site-prepared and planted with 

these other southern pines. Restoration requires removal 

of the loblolly or slash pine overstory and establishment 

of longleaf pine seedlings. If possible, prescribed burning 

should be conducted ~ 2 years prior to the harvest, to reduce 

woody competition and stimulate growth of herbaceous 

understory species. A site-preparation fire following logging 

is needed to remove debris and discourage hardwood trees 

and shrubs. Chopping may be used after harvesting and 

prior to the site-preparation fire to help control woody plant 

competition and reduce the density of loblolly and slash 

pine seedlings. A single-drum chopper should be used to 

avoid excessive soil disturbance. Repeated prescribed fires 

after establishment will determine which species are best 

adapted to particular microsites. Although longleaf pine 

once dominated these sites, loblolly and slash pines often 

occurred with them as natural components of these forests. 

Therefore, the objective is not to totally eliminate competing 

pines, but rather to provide longleaf pine with an advantage 

that compensates for past actions that selected against it. 

Understories on these wetter sites seem to be more resistant 

to changes caused by soil disturbance, probably because 

moisture availability is greater on such sites than on the sandhills (Outcalt and Lewis 1990). Therefore, many existing 

slash and loblolly pine plantations have fairly complete and

healthy understory communities, even if the sites were bedded

when the plantation was established. As a consequence, 

some managers prescribe bedding prior to planting of bareroot or containerized longleaf pine seedlings for restoration, 

to increase survival rates. However, extensive studies indicate

that survival of slash pine is not increased by this reduction 

in competition. Even where bedding was done following 

double disking, a practice that greatly reduces competition, 

slash pine survival after 10 years was not improved over that 

on wet flatwoods sites in northern Florida that were only 

burned (Outcalt 1983). The improved microsite conditions 
21

created by bedding do increase survival rates in years that are

wetter than normal. However, on average, the increase in slash

pine seedling survival is only ~ 15 percent (Schultz 1976). 

Although adult longleaf pines grow on very wet sites, longleaf pine seedlings seem more susceptible to mortality than

slash pine seedlings during periods of soil saturation (Wahlenberg 1946). Under natural conditions, longleaf pines very 

likely become established on wet sites during drier periods. 

Bedding could be expected to increase longleaf pine seedling survival rates during wetter years by > 15 percent. However, this survival gain must be weighed against the extra 

cost of bedding and against the damage to the native groundcover. Bedding may also alter site moisture relations and 

nutrient distribution for > 30 years (Schultz 1976). In such a

case, the ecosystem will not be truly restored even if longleaf

pine is successfully established and native species dominate 

the understory. Restoration methods can be modified on wet 

sites to improve survival without resorting to bedding. It 

would be more economical and ecologically advantageous to

plant additional longleaf pine seedlings, as a hedge against

lower survival, than to bed a site. Other adjustments can be

made: for example, planting can be done during drier seasons

or postponed when sites are flooded and longleaf pine seedling survival would likely be low. Although container seedlings could be planted where this is justified on economic 

or other grounds, it is also reasonable to accept lower rates 

of longleaf pine survival or to accept a resulting mixture of 

longleaf pine and other southern pines that naturally occur 

in the wetter depressions embedded in flatwoods areas. 

Upland and montane sites dominated by longleaf pine 

with native understory—Few upland and mountain sites 

remain in longleaf pine, because these sites were preferred 

for agricultural, urban, and residential development. However, many of the longleaf pine forests that now exist on 

such sites are in fair to good condition. These include sizable

areas on Fort Bragg in North Carolina, which have been 

subjected to frequent fires associated with military training 

activities. A substantial area of upland longleaf pine ecosystem exists on large hunting plantations in the Red Hills area

of southwestern Georgia and is maintained by frequent burning. Relatively healthy montane longleaf pine forest still 

exists at Fort McClellan in Alabama, where frequent fires 

associated with military training activities have maintained 

the open understory and discouraged invasion by woody species (Varner and others 2001). However, upland areas mostly 

in Alabama, Mississippi, Louisiana, and Texas and montane 

sites in Alabama and Georgia have developed unnaturally 

dense hardwood midstories and require restoration. Because 

these are among the most biologically productive longleaf 

pine sites, they change most rapidly, quickly developing

midstory layers in the absence of frequent fire. In addition 

to a very dense midstory and a shrub-dominated understory, 

these sites also accumulate potentially hazardous quantities 

of fuel. Frequent growing-season fires are needed to adequately control competing woody plants on upland sites 

with better soils. As on flatwoods sites, frequent growingseason fires over many years are required to reduce the hardwood rootstocks (Boyer 1990a). As in other longleaf pine 

ecosystem types, a series of dormant-season fires may be 

necessary to gradually reduce fuel levels before growingseason burning begins. 

In a common variant of this ecosystem type, longleaf pine

is present, but other southern pines are also present as dominants or co-dominants. In addition to prescribed burning as

outlined above, these stands need selective harvesting to 

reduce the presence of other southern pines and hardwoods 

in the overstory. The objective is not total elimination of 

other tree species, but rather a readjustment of overstory composition, recognizing that these other species are part of the 

natural longleaf pine community. Burning should commence 

before selective harvesting begins, to prevent woody plants 

from proliferating and forming a shrub thicket in openings 

that will be created by harvesting operations. Herbicide application and mechanical reduction of nonmerchantable woody 

species may also be useful in accelerating the process of 

readjusting species composition and dominance (Boyer 1991). 

Upland and montane sites dominated by other species—

Little research information or management experience is 

available to guide restoration on these sites. Most native longleaf pine forests on such sites were converted to agriculture

or loblolly pine plantations, and this conversion significantly 

altered composition of the understory plant community. The 

few sites that show no evidence of severe soil disturbance 

support scattered natural longleaf pine trees in a mixture 

dominated by loblolly pine, shortleaf pine, and hardwoods. 

Restoration is made more difficult by the high biological

productivity of these sites. Repeated and prolonged treatment

with prescribed fire should eventually reduce the abundance 

and cover of woody plants in the understory. It is probable 

that at least a portion of the native understory still exists in 

the soil seed bank or as suppressed individuals (Varner

and others 2000). Therefore, restoration would begin with 

burning to reduce fuel and initiate control of woody shrubs 

and hardwoods. If timber markets allow, selective harvesting 

could then be used to release any native longleaf pine and 

reduce the hardwood component. Otherwise, thinning would 

be performed at a financial cost. Other pines may need to be 

retained onsite to furnish sufficient needle fall for prescribed 

burning and to prevent release of woody competition. Once

prescribed burning and other mechanical or chemical methods
22

have reduced the woody midstory and understory layers, 

some of these other pines could be removed and replaced 

with longleaf pine seedlings. The best way to do this would 

probably be to create canopy gaps in areas where the understory has become dominated by grasses and forbs. 

Development of restoration techniques for upland sites once 

used for agriculture or intensive forestry has begun only 

recently. It is unlikely that many of the native understory 

grasses and forbs survived intensive soil disturbance; however, there is a large soil seed bank of herbaceous weeds that

must be controlled. A restoration technique being tested consists of multiple-pass harrowing to reduce weeds followed 

by direct seeding of native species. Although native species 

have been established by this method, their long-term survival

and growth are as yet uncertain.2 There is a need to develop 

new techniques that would provide the option to restore a

more complete biotic community on the many areas of privately owned land that have been reforested with longleaf 

pine.

Restoration Efforts

Restoration of longleaf pine forest ecosystems first began on

public lands as part of an effort to halt decline of the endangered red-cockaded woodpecker. Initial efforts focused on

mechanical removal of midstory hardwoods near cavity nest

trees and was augmented with prescribed fire to control hardwood sprouts. This effort later expanded to a more general 

approach of improving habitat for all species, with special 

consideration given to imperiled species. As a major public 

landholder, the USDA Forest Service conducts an active 

program for restoring longleaf pine ecosystems on national 

forests in the Southeast. In the late 1990s, longleaf pine 

ecosystems occupied 307 000 ha on national forestlands 

(McMahon and others 1998). To improve the condition of

these forests, managers have begun to implement more 

burning during the growing season on a 3- to 4-year cycle. 

Mechanical removal and herbicide treatments are used, 

where appropriate, to augment fire. Loblolly pine and slash

pine are being removed, and longleaf pine is being reestablished on its former sites. The goal is to increase the area 

occupied by longleaf pine ecosystems by 50 percent, to a 

total of 451 600 ha. 

The U.S. Department of Defense also has substantial land 

holdings within the native range of longleaf pine. One of 

the largest of these holdings, Eglin Air Force Base, is an 

important part of the largest remaining concentration of longleaf pine habitat, the gulf coast area of Florida and Alabama 

(Outcalt and Sheffield 1996). This base contains large areas

of the longleaf pine sandhills ecosystem that has been maintained in healthy condition by the frequent fires resulting 

from military training activities. However, it also contained a 

large area of forest with a well-developed midstory of scrub 

oaks. In 1993, land managers began an ambitious restoration 

program that employed fire and supplemental mechanical 

treatments and herbicides when these were needed. Between 

1993 and 1997, they burned an average of 16 200 ha/year, 

with 70 percent of these prescribed fires being conducted 

during the growing season (McWhite and others 1999). This

significantly reduced the hardwood midstory and improved 

the composition and density of the understory herbaceous 

plant community. Managers also selectively harvested sand 

pine from 4750 ha of longleaf pine stands it had invaded 

during the period of fire exclusion. Between 1993 and 1997,

offsite slash pine (1360 ha) and sand pine (1950 ha) plantations were harvested and replaced with longleaf pine seedlings. Manual planting was used on all sites, and 90 percent 

of the seedlings were containerized to improve survival. A 

smaller area of coastal longleaf pine flatwoods at Eglin Air 

Force Base is being restored through the reintroduction of 

prescribed fire. 

Other military bases across the region have been following 

or are adopting similar policies and practices. Fire is being 

reintroduced where needed, and areas occupied by offsite 

species are being converted back to longleaf pine, with 

recognition that the understory plant community must be 

conserved during this process. Military installations on 

the lower Coastal Plain, such as Camp Lejeune in North 

Carolina and Fort Stewart in Georgia, have many areas of 

flatwoods and lowland longleaf pine ecosystems. Substantial 

restoration is taking place in these wetter longleaf pine 

ecosystems as well as in the drier sandhills. 

Restoration activities are taking place on almost all other 

Federal and State lands that have longleaf pine or sites suitable for its establishment. Private landowners with larger

tracts have used periodic prescribed fire in their management, and there is growing interest in doing at least a portion

of this burning during the growing season. However, because

of the high cost of treating small parcels, most small landowners are unable to burn their sites (Outcalt 1998), and 

very little restoration is occurring on these areas.

Understory restoration on areas that have been substantially 

altered is more difficult and expensive. Therefore, it is not 

surprising that only a limited amount of restoration work is 

being done on such sites. Wiregrass plugs are being grown

2 Personal communication. 2002. L. Katherine Kirkman, Research Scientist,

J.W. Jones Ecological Research Center, Rt. 2, Box 2324, Newton, GA 

31770.
23

and outplanted along with longleaf pine seedlings on both

State land (Pittman and Karrfalt 2000) and national forestland in Florida. At the Savannah River Site in South Carolina,

the USDA Forest Service has recently planted wiregrass 

plugs under longleaf pines planted on old-field sites during 

the 1950s, to restore fuel and improve the habitat of redcocked woodpecker colony sites. The Nature Conservancy is 

restoring both longleaf pine and wiregrass on a sandhills site 

at their Apalachicola Bluffs and Ravines Reserve in northern 

Florida (Hattenbach and others 1998, Seamon 1998). They 

are also experimenting with direct seeding of other native 

understory species. The most extensive program for direct 

seeding of understory species is being conducted at Fort 

Stewart (table 4). Resource managers have collected seed 

and sown it on site-prepared areas since 1997.3 A tractormounted flail-vac seed harvester is used to collect seed from 

onsite areas burned during the growing season. Collections 

vary from 750 to 1100 kg/year, which at a mean sowing 

rate of 13.2 kg/ha is enough to sow 57 to 83 ha/year. Seed is 

spread on restoration sites by means of a platform-mounted 

bale chopper on the back of a farm tractor. Flatwoods and 

sandhills sites are being restored with seed from appropriate 

donor sites. The goal at Fort Stewart is to restore 8100 ha of 

old fields to functioning longleaf pine ecosystems (Hilliard 

1998).

Costs and Benefits Associated 

with Restoration

Estimating Restoration Costs

Disagreement over what constitutes restoration and lack of

precise criteria for judging the success of restoration make

estimating the cost of restoring longleaf pine forest ecosystems

quite difficult. If the focus is limited to restoring longleaf 

pine as the dominant overstory tree through reforestation, 

then reasonable cost estimates can be made using readily 

available economic data. However, if the intent is to restore 

a completely functional ecosystem, then cost estimation 

becomes very difficult. The necessarily long-term nature of

ecosystem restoration also introduces temporal considerations that significantly affect costs incurred by forest managers. Although this time frame can be shortened somewhat 

by management techniques that accelerate the restoration 

process, the use of these techniques may involve increases in 

short-term costs. 

Reestablishing longleaf pine as the dominant tree species on 

a site is often the first and, in many ways, easiest step in the

restoration process. Reforestation costs vary according to

ambient conditions and the type and amount of site preparation

needed to achieve successful establishment of tree seedlings. 

On previously harvested or old-field sites, costs typically 

range from $370 to $740/ha, depending on site conditions 

and whether bare-root seedlings or containerized seedlings 

3 Personal communication. 2001. Dena Thompson, Wildlife Biologist, 

Department of Environmental Health, Forestry Branch, Fort Stewart, GA 

31314.

Table 4—Production rates, equipment, and costs for understory plant restoration at Fort Stewart, GA

Seed collectiona Seed sowing

Year Days ha kg kg/day kg/ha Days ha ha/day kg/ha

1998 17 61.9 821 48.3 13.3 12 62.3 5.2 13.2 

1999 24 79.5 938 39.1 11.8 14 74.1 5.3 12.7 

2000 20 20.2 539 27.0 26.7 9 40.5 4.5 13.3 

2001 30 95.1 1096 36.5 11.5 16 97.2 6.1 11.3 

2002 17 57.9 746 43.9 12.9 6 40.5 6.8 18.4 

Mean 22 62.9 828 37.6 13.2 11 62.9 5.7 13.2 

4-wheel-drive vehicle $47,594.00 Seed collection labor $10.74/kg 

Flail-vac seed collector $11,950.00 Sowing labor $58.05/ha 

Small tractor $35,000.00 Seed costb

 $141.77/ha 

Bale chopper $3,800.00 Understory restoration costb $199.82/ha

a Seed collection season varies from late October to late December.

b Does not include equipment purchase or operation and maintenance costs.
24

are selected. This range reflects the current costs for site 

preparation, seedlings, and planting. To control competing 

vegetation, increase survival, and stimulate early growth, an

additional $85 to $100/ha might be expended for herbicide 

application. Because these costs occur early in the investment,

they cannot be discounted over time when calculating net 

present value, internal rate of return, or other economic 

indices. Despite these expenditures, the average internal rate 

of return for such an investment is estimated at 10.1 percent, 

with a range from 8 to 12 percent (Busby and others 1996). 

On sites that already support forests, the presence and distribution of longleaf pine in the canopy may be sufficient to 

allow for natural regeneration. In such cases, unwanted tree

species should be harvested and removed, leaving the longleaf pine as the dominant seed-producing tree for future 

generations. Prescribed fire or selective herbicides or both 

are usually required to prevent woody shrubs and trees from 

developing into a midstory layer and to prepare a seedbed of 

mineral soil favorable for germination of longleaf pine seed. 

While natural regeneration appears to be free, determining 

its true cost in economic terms requires assessing assumed 

risk, calculating opportunity cost, and considering the timevalue of money. 

In forests with degraded understories or understories composed

of nonnative plants, establishing longleaf pine as the dominant overstory tree is only the beginning of the restoration 

process. Without the species richness and diversity unique to 

their native plant and animal communities, these ecosystems 

cannot be considered fully restored. Restoring groundcover 

plants can be very expensive, with costs rising sharply as the 

time scale is compressed. In relatively undisturbed forests, 

many plants native to the site may be reestablished by the 

reintroduction of fire, particularly growing-season fire. Fires 

facilitate reestablishment of vascular plants by stimulating 

seed banks and inducing flowering and seed production in 

many existing plants. The cost of fire reintroduction varies 

with existing site conditions. Long-term fire exclusion can 

cause fuel accumulations that make fire reintroduction potentially very destructive. Mitigating this danger can be quite 

labor intensive and very expensive. In frequently burned 

areas, the risks and cost of burning are greatly reduced. In

such cases, changing the fire frequency or season of burning 

or both may be sufficient to initiate the recovery of native 

plant communities. Where seed banks are depleted or severe

soil disturbance has occurred, restoring the plant community

is more problematic. Reseeding or replanting selected understory plant species has been accomplished successfully, but

at considerable economic cost. Estimates of the cost of artificially reestablishing sustainable populations of native 

groundcover plants range from several hundred to several 

thousand dollars per hectare. Techniques for seed collection, 

cultivation, distribution, planting, and other steps in the process are only in the developmental stages at this time and are

generally focused on pyrophytic graminoids (e.g., wiregrass),

species consumed by wildlife (e.g., legumes), and species 

of special concern (e.g., American chaffseed, Schwalbea 

americana). 

Restoration of a diverse understory of native plants and a

suitably structured overstory are viewed as crucial in reestablishing and sustaining viable populations of the many wildlife

species indigenous to these ecosystems. At this time, efforts 

to restock these animals are very limited, because the costs

of such programs are high and because capture and relocation permits must be obtained from State and Federal authorities. Efforts to restock pocket gophers and fox squirrels, both

indicators of healthy ecosystems, are virtually unknown and

would likely be unsuccessful at any cost if the plant community does not provide suitable habitat conditions. Because 

northern bobwhite are game birds of substantial economic

importance and are well adapted to longleaf pine ecosystems,

their establishment and maintenance has received significant 

attention. The red-cockaded woodpecker is a major indicator

of the health of longleaf pine ecosystems, and costs associated with its recovery represent a substantial component of

the cost of restoring these ecosystems. Red-cockaded woodpeckers have exacting habitat requirements. They require a 

relatively open overstory canopy with widely spaced mature 

pines, trees of large diameter that may serve as nesting sites 

following cavity excavation, a sparse or absent midstory, 

and groundcover vegetation that is low in stature. Wildlife 

biologists consider the creation of these conditions as being 

synonymous with longleaf pine ecosystem restoration. 

Restoring red-cockaded woodpeckers to areas of suitable 

but unoccupied habitat is a highly regulated and expensive 

activity. Costs of such efforts include (1) construction of 

artificial cavities in large pines; (2) capture, transportation, 

and release of the birds; and (3) population monitoring for 

an extended period after relocation. 

Benefits of Restored Longleaf Pine Ecosystems

Both the material and intangible benefits resulting from 

restoration of longleaf pine ecosystems are substantial. The 

economic value of longleaf pine forests is considerable, and

commercial products can be extracted from a properly functioning forest without significantly disrupting ecological 

processes. Longleaf pine is the most versatile of all the 

southern pines and provides a wide variety of products, many

of which are highly valued (table 5). Longleaf pine forests 

typically produce up to five times more tree stems of sufficient quality to be used as utility poles than do stands of

slash pine or loblolly pine. Stumpage values for such poles 
25

exceed prices for sawtimber by ~ 40 percent. When the high

value of pine straw (i.e., fallen needles, which may be harvested from stands as early as age 10) is added, the economic

value of longleaf pine forests becomes increasingly obvious. 

Contingency or “willingness to pay” surveys consistently 

indicate that hunter access to private lands has value as a 

tradable commodity throughout the natural range of longleaf 

pine. Where longleaf pine forests are maintained in open 

parklike condition, the higher quality of this habitat for 

quail, turkey, and deer brings premium economic returns 

in the form of hunting leases and related services to private 

landowners. Revenues correspondingly decline as habitat 

conditions become degraded.

Enhanced recreational opportunities, greater esthetic appeal, 

and higher real estate values associated with restored longleaf pine ecosystems are attributes that are more difficult to

measure. However, ecological values resulting from restoration of ecosystem health are perhaps the greatest benefits 

that defy easy quantification (Constanza and others 1992). 

Redevelopment of appropriate community structure and

composition and reinitiation of essential ecological processes promote improved ecosystem resilience and stability. 

Ecosystem services are of inestimable value and include 

climate amelioration, water purification, soil stabilization, 

flood control, nutrient cycling, providing habitat, and serving as a reservoir for genetic material that could be a source 

for new medicines, energy, and industrial feedstock (Burton 

and others 1992, Ledig 1988, Riggs 1990).

Table 5—Value of wood products from major southern 

pine speciesa

Sawtimber High-quality 

price/m3 poles Value/ha

Longleaf pine $264 66% $8,492

Slash pine $257 12% $7,640

Loblolly pine $254 5% $7,454

a Assuming a mean stand volume of ~ 29 m3

/ha at age 55 (Holliday 2001).

Influence of Forestland Ownership

Regional Patterns

Of the 87 million ha of forest in this region, 81.3 million 

ha are classified as timberland, that is, land capable of 

producing forest products and not withdrawn from potential timber harvesting for special uses (Conner and Hartsell 

2002). Timberland in the region is held by many groups, 

which can be broadly divided into public and private ownerships (table 6). About 8.7 million ha of this timberland is 

publicly owned, including lands administered by Federal, 

State, county, and municipal government agencies, as well 

as by the tribal governments of Native Americans. The USDA

Forest Service manages the largest area of public land, with 

4.7 million ha under its administration. Private interests own 

the remaining 72.6 million ha of timberland in the region. 

Forest industry manages 15.1 million ha, other corporations 

own 8.8 million ha, and nonindustrial private landowners hold

48.7 million ha. It is worth noting that the total area owned by

private individuals has increased by 18 percent since 1982. 

One potential result of this increase is forest fragmentation, 

the division of larger contiguous tracts of forest into smaller, 

more isolated parcels. Although it was not monitored closely 

in the past, information on the tract size of forestland is now 

being collected to aid in the forecasting of future trends in 

ownership and landscape condition.

Table 6—Regional forestland ownership

 Percent of

 Million ha forestland

Forestland 87.0 100.0

Nontimberland 5.7 6.6

Timberland 81.3 93.4

 Public 8.7 10.0

 USDA Forest Service 4.7 5.4

 Other Federal agencies 1.9 2.2

 State and local agencies 2.1 2.4

 Private 72.6 83.4

 Forest industry 15.1 17.3

 Other corporations 8.8 10.0

 Nonindustrial private 48.7 56.0

 landowners

Source: Conner and Hartsell (2002).
26

Land Ownership Patterns in Longleaf 

Pine Ecosystems

There are now < 1.2 million ha of longleaf pine forest in the

Southeastern United States, and much of that is in poor condition. About 51 percent of the existing longleaf pine forest

is owned by nonindustrial private landowners, 16 percent by 

forest industry, and 33 percent by public land management 

agencies (Outcalt and Sheffield 1996). Public lands constitute only 10 percent of forestland in the region, but they 

support a larger percentage of the area in longleaf pine. 

Ownership stability over the long term and public agency 

ecosystem management programs that do not exclusively 

emphasize commodity production provide a more secure 

habitat for this long-lived tree species that can be sustained 

by less intensive management practices. Public lands also

more often exist as larger, less fragmented tracts with linkages that provide ecological connections among otherwise 

isolated longleaf pine forest “islands.” The many resource 

values and desirable ecological attributes of longleaf pine 

forests also complement the land management mission of 

most public agencies. While immediate emphasis should be 

placed on restoring lands currently occupied by degraded 

longleaf pine forests, estimates indicate that several million 

additional hectares may ultimately be suitable for restoration 

to longleaf pine ecosystems. 

A recent survey of longleaf pine stands in North Carolina, 

South Carolina, Georgia, and Florida revealed that ~ 50 

percent of the sites currently dominated by longleaf pine are 

in fair to good condition with a healthy understory of native 

plant species, no extensive midstory, and little or minimal 

soil disturbance (Outcalt 2001). If the western portion of the 

range has a similar distribution of conditions, the remaining 

area of healthy longleaf pine ecosystems is ~ 560 000 ha. An 

additional 480 000 ha is also dominated by longleaf pine but 

requires restoration to reduce the midstory layer or improve 

the understory plant community or both. Most of the latter 

areas (400 000 ha) are owned by private nonindustrial forest

landowners, while the 80 000 ha on public lands have not 

yet been restored. A 1995 survey found another 180 000 ha

of longleaf pine growing as plantations on lands owned by

forest industry and nonindustrial private landowners. Since

that time, an additional 70 000 ha of longleaf pine plantations

have been planted on private lands with funding provided by

the CRP. Most of these sites have been altered greatly (especially the CRP areas), and it would require considerable 

investment to fully restore native longleaf pine ecosystems 

on them. It is estimated that public agencies other than the 

USDA Forest Service control about 200 000 ha of lands that

have some native understory components and are dominated 

by other overstory tree species (McMahon and others 1998).

USDA Forest Service inventory and analysis data indicate 

that > 600 000 ha of privately owned forestland that is dominated by other tree species (mostly scrubs oaks, loblolly 

pine, or slash pine) supports at least some of the understory 

plants native to longleaf pine ecosystems. This land is potentially available for restoration by replacing the current overstory dominants with longleaf pine. There are additional 

plantations of other tree species across the entire former 

range of longleaf pine that could be restored to the longleaf 

pine ecosystem, but for a majority of these, this would be a

difficult task. Overall, we estimate that there are 280 000 ha

on public lands and 1 million ha on nonindustrial private 

lands that could be fully restored at a reasonable cost with 

current knowledge. Clearly, the greatest opportunity for 

future restoration of longleaf pine ecosystems resides with 

nonindustrial private forest landowners.

Influence of Ownership on Restoration

Individual forest landowners elect or decline to restore longleaf pine forest ecosystems on suitable lands for a variety of 

reasons. Private nonindustrial landowners are diverse and 

have widely varying land management objectives. Generally, 

the strongest incentives for private nonindustrial landowners 

to proceed with restoration activities are government-funded 

programs that share the cost of artificially regenerating and 

maintaining longleaf pine forests. Without these cost-share 

programs, many landowners would not consider longleaf 

pine restoration as a land management option, because any 

short-term financial reward for doing so would be absent. 

However, some affluent landowners do implement longleaf 

pine restoration on their lands without seeking financial 

assistance from government programs. They do so because 

they feel it is the ecologically right thing to do or because 

they wish to leave a natural heritage legacy to their heirs. 

Forest industry and other corporations in the region typically 

manage forests on short rotations (< 30 years) for fiber products in order to maximize profits, especially in the short 

term. Because longleaf pine must usually be grown on longer

rotations to produce larger trees for sawtimber, poles, and 

pilings, forest industry has generally shown less interest in 

restoring and managing longleaf pine forest ecosystems than 

have other landowners. Some private industrial companies 

do infrequently grow longleaf pine forests on their lands in 

an attempt to diversify their financial assets and contribute 

to the achievement of broader resource conservation goals. 

During the past several years, some companies have begun 

establishing longleaf pine on sites that are better suited to 

it than to loblolly pine or slash pine, and at least one forest 

industry firm is planning an ambitious program to restore 

longleaf pine forests across a substantial area. 
27

In recent years, numerous public land management agencies

have actively sought to restore longleaf pine forest ecosystems on national and State forests, national and State wildlife refuges, State parks, military bases, and other public 

lands. Longleaf pine restoration efforts undertaken in accordance with ecosystem management policies are a major component of the long-range mission of various public agencies 

to protect populations of rare plants and animals, conserve 

biological diversity, and provide the public with multiple 

benefits such as recreational opportunities, clean water, wood,

and other forest products. Unfortunately, costs associated 

with restoration efforts and the regulatory and procedural 

constraints on forest management practices that are necessary to convert sites occupied by other forest types to longleaf pine forests have impeded the rate of restoration of 

longleaf pine ecosystems on public lands.

Research and Management Needs and 

Policy Implications

Scientific Knowledge

For decades, southern research and development efforts by 

industry, government, and universities focused on shortrotation plantations of loblolly pine and slash pine for paper 

pulp production, with little investment in research work on 

longleaf pine regeneration, growth, and productivity (Boring 

1999). Although increasing research emphasis on longleaf 

pine in recent years has produced some very encouraging 

results, a great many questions remain partly or largely unanswered. An entire publication would be required to present 

an exhaustive list of longleaf pine research needs (Walker 

1995). Therefore, only major topics of scientific inquiry are 

summarized below.

Although they are incomplete, historical data about the composition, structure, and function of natural ecosystems may

serve as a reference to promote general understanding (White

and Walker 1997). These include quantitative and qualitative 

descriptions of (1) understory components and the processes

by which they were established and maintained, (2) age and

spatial structure of even-aged and uneven-aged forests and

the processes that influenced canopy condition, (3) landscape

structure and its variation across the region, and (4) the interaction between fire regimes and landscape structure, particularly as this interaction influenced ecosystem processes 

(Walker and Boyer 1993). Better quantitative data about 

existing longleaf pine forest ecosystems would facilitate comparative analyses that would support appropriate restoration 

efforts. Such analyses include (1) examination of local biotic 

community quality and degree of deviation from historical 

patterns; (2) assessment of the responses of ecosystems in 

various conditions to restoration treatments and fire regimes; 

(3) identifying and explaining the processes that underlie the

population dynamics of native plants and the influence of

silvicultural treatments on them; (4) quantitatively describing landscape scale structures, flows, and functions and 

showing how these currently depart from historical patterns; 

and (5) cumulative effects evaluation of historical land 

uses, contemporary management treatment regimes, and 

catastrophic events such as hurricanes and fires. 

Because ecological restoration is a management process that

is essentially goal-driven, it is essential that the desired result

be defined as precisely as possible. Various kinds of basic 

knowledge can help us identify realistically appropriate 

target conditions. Such scientific knowledge includes (1) 

patterns of genetic variation in longleaf pine and associated 

native species and the effects of habitat fragmentation on 

this variation, (2) life-cycle dynamics of rare plant and 

animal species and the factors essential for population viability, (3) the forest site and microsite affinities of native 

species, (4) the influence of natural and anthropogenic disturbances at local and landscape scales and especially the

influence of fire as a driving ecological process, (5) ecological thresholds related to multiple stable states and how they 

are influenced by various disturbances, and (6) appropriate 

criteria for defining longleaf pine ecosystem health and 

measures of ecosystem integrity that can be monitored 

effectively. 

Management Information

Forestland managers involved in restoration efforts throughout the region have identified many pressing management 

information needs. Key among these needs is a better understanding of the ecology and dynamics of fire in longleaf pine 

forests, including (1) the effects of growing-season fire and 

dormant-season burning in a wide range of habitat types; 

(2) appropriate fire frequencies, intensities, and ignition 

techniques to achieve various objectives; (3) the influence 

of fire on important game species and numerous nongame 

species; (4) the role of fire in sustaining various at-risk 

species and the selection of burning methods to ensure their 

continuing viability; (5) techniques for safely reintroducing 

fire into long-unburned ecosystems and mitigating loss of

resource values; (6) the degree to which and under what conditions chemical and mechanical treatments can be substituted 

for periodic prescribed fire (especially at the wildland-urban 

interface, where burning may be restricted); and (7) models 

that will enable managers to better predict and manage 

smoke resulting from prescribed burning. 
28

Successful regeneration of longleaf pine is an essential 

element of efforts to restore and sustain these ecosystems. 

Information needs in this area include improved understanding of (1) the structures and processes that affect longleaf 

pine regeneration, including canopy-gap and root-gap dynamics and the influences of light, nutrients, moisture, and fuels

that support fire on seedling survival and growth; (2) variation in intraspecific and interspecific competition across a

wide range of site types; (3) silvicultural methods for effectively regenerating longleaf pine in uneven-aged, two-aged, 

and even-aged forests and the use of these to mimic natural 

disturbance patterns; and (4) appropriate densities for planting longleaf pine seedlings across a range of site types.

Conservation efforts that do not consider social and economic

costs and benefits are usually doomed to fail in the long term

(Kimmins 1992, Oliver 1992). To be successful, restoration 

efforts aimed at sustainable management must provide opportunities to achieve stewardship goals and allow for the use 

of natural resources. Information needed to address these 

concerns includes (1) periodically updated estimates of the 

economic costs and expected returns of restoring longleaf 

pine, (2) accurate projections of the quantity and quality of

longleaf pine wood products (and pine straw, where appropriate) that can be expected to result from restoration efforts, 

(3) economical methods for reestablishing both overstory trees

and understory plants, (4) impacts of logging and replanting

on longleaf pine forest environments, and (5) public knowledge about and attitudes toward longleaf pine ecosystem 

restoration, conservation, and utilization.

Restoration of longleaf pine forests requires looking beyond 

the trees and considering, to the greatest degree practical, all

ecosystem components. In striving toward this goal, managers

need information on (1) the effectiveness of various mechanical, chemical, and pyric site preparation techniques and their

impacts on native plants in the understory; (2) techniques for 

efficiently establishing native understory plants; (3) methods 

for suppressing invasive species and discouraging the spread 

of nonnative species; (4) methods for restoring forests across

the full range of suitable site types and techniques for improving degraded longleaf pine ecosystems; and (5) computerized

management support systems (i.e., expert systems) that 

assist managers in identifying potential problems and opportunities that arise during restoration efforts.

Policy Implications

Public policies promoted by government agencies that facilitate stewardship and restoration of longleaf pine ecosystems 

have long been in need of implementation (Boring 1999). 

During the period when management of longleaf pine was

discouraged by Federal and State foresters, it seemed reasonable that little if any public funding should be expended to

support programs for this species. However, social attitudes 

toward longleaf pine forests have changed in recent decades, 

with interest in these ecosystems rising each year. In response

to this change, greater public funding could be allocated for 

programs that develop and disseminate new knowledge in 

support of managers interested in longleaf pine. 

As ecosystem management has become the dominant theme 

on public lands, goals for restoring longleaf pine have been 

established throughout the region. There are indications that

the area in longleaf pine ecosystems could be doubled in the

future. However, such an effort would require the endorsement of policymakers and need substantial financial support 

over a long period. Although longleaf pine restoration efforts

on private lands have been impeded by various factors, a frequently lacking ingredient for success is timely and appropriate management information for nonindustrial private 

landowners. Clearly, greater resources are needed to provide 

technical assistance to these important members of the 

southern forestry community. 

Conservation and restoration of longleaf pine ecosystems 

might be more effectively achieved by creating a reserve 

program modeled on the Wetlands Reserve Program, which 

has been very successful in maintaining wetland habitats 

at the State and national levels. Longleaf pine ecosystems 

have declined by 97 percent, while the historical losses 

of wetlands are estimated at only 47 percent. Certainly, 

longleaf pine ecosystems deserve such consideration.

Many private landowners are seriously concerned about the

implications of managing forests that also may serve as 

habitat for endangered or threatened species. Technical assistance programs could not only provide important habitat 

management guidance, but also advise landowners on how

to enroll in Safe Harbor and conservation easement programs

to effectively address these concerns (Costa 1999). Also, the

need to periodically burn longleaf pine forests and the limitations placed on prescribed fire by government regulations 

have dissuaded many landowners from managing longleaf 

pine on their lands. Right-to-burn laws would, to a substantial degree, address this issue and need to be adopted widely 

in the region.

Urbanization at the wildland-urban interface continues to 

result in loss of longleaf pine sites. Such urban development 

fragments land ownership into a series of smaller tracts that

are permanently lost from the land base of functioning longleaf pine ecosystems. The current structure of estate and 

property tax laws also places a heavy financial burden on 
29

private landowners, with the result that large land holdings 

are frequently divided and sold as smaller parcels, further 

fragmenting the ownership of longleaf pine lands. Although 

use of conservation easements tied to land deeds affords tax

relief while diminishing fragmentation, and recent tax law

reform provides some benefits to landowners, these developments have been insufficient to change this negative trend. 

The substantial initial costs of establishing longleaf pine can

also be a problem for many landowners, despite the fact that

several of the government assistance programs discussed 

earlier aid in partially offsetting some of these costs. Overall,

financial difficulties still remain a major disincentive for 

many landowners. Therefore, perhaps the greatest policy 

need for the private sector is a comprehensive change in tax

laws to provide specific and substantial benefits to landowners

who restore and maintain longleaf pine forest ecosystems on 

their lands. 

Acknowledgments

The authors are grateful to Becky Estes for searching the

literature to identify numerous relevant publications, and

to Dave Haywood, Steve Jack, John Kush, Dave Borland,

Ric Jeffers, John Stanturf, Palle Madsen, Alan Salmon,

Paul Smith, and Louise Wilde for comments helpful in 

improving this manuscript.

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Department of Agriculture, Forest Service, Southern Research Station. 34 p.

Longleaf pine (Pinus palustris) ecosystems once occupied 38 million ha in the Southeastern 

United States, occurring as forests, woodlands, and savannas on a variety of sites ranging from 

wet flatwoods to xeric sandhills and rocky mountainous ridges. Characterized by an open parklike 

structure, longleaf pine ecosystems are a product of frequent fires, facilitated by the presence 

of fallen pine needles and bunchgrasses in the understory. Timber harvest, land conversion to 

agricultural and other nonforest uses, and alteration of fire regimes greatly reduced longleaf pine 

ecosystems, until only 1.2 million ha remained in 1995. Longleaf pine ecosystems are among 

the most species-rich ecosystems outside the tropics. However, habitat loss and degradation 

have caused increased rarity of many obligate species. The lack of frequent surface fires and the 

proliferation of woody plants in the understory and midstory have greatly increased the risk of 

additional longleaf pine ecosystem losses from catastrophic fire.

Because longleaf pine still exists in numerous small fragments throughout its range, it is reasonable to conclude that it can be restored. Restoration efforts now underway use physical, chemical, 

and pyric methods to reestablish the natural structure and function in these ecosystems by adjusting

species composition, modifying stand structure, and facilitating ecological processes, such 

as periodic fire and longleaf pine regeneration. The ecological, economic, and social benefits 

of restoring longleaf pine ecosystems include (1) expanding the habitat available to aid in the 

recovery of numerous imperiled species, (2) improving habitat quality for many wildlife species, 

(3) producing greater amounts of high-quality longleaf pine timber products, (4) increasing the 

production of pine straw, (5) providing new recreational opportunities, (6) preserving natural and 

cultural legacies, and (7) creating a broader range of management options for future generations.

Keywords: Biological diversity, bluestem grasses, disturbance, fire ecology, gopher tortoise, Pinus 

palustris Mill., red-cockaded woodpecker, wiregrass. 

METRIC CONVERSION TABLE

Symbol When You Know Multiply by To Find Symbol

LENGTH

 mm millimeters 0.0394 inches in

 cm centimeters 0.3937 inches in

 m meters 3.281 feet ft

 m meters 1.094 yards yd

 km kilometers 0.6214 miles mi

AREA

 cm2 square centimeters 0.1550 square inches in2

m2 square meters 1.196 square yards yd2

 km2 square kilometers 0.3861 square miles mi2

 ha hectares 2.471 acres 

 (10 000 m2)

MASS (weight)

 g grams 0.0353 ounces oz

 kg kilograms 2.205 pounds lb

 t metric ton 1.102 short tons 

 (1 000 kg)

VOLUME

 mL milliliters 0.0338 fluid ounces fl oz

 mL milliliters 0.0610 cubic inches in3

 L liters 2.113 pints pt

 L liters 1.057 quarts qt

 L liters 0.2642 gallons gal

 m3

 cubic meters 35.315 cubic feet ft3

 m3 cubic meters 1.308 cubic yards yd3

TEMPERATURE

oC degrees multiply by 9/5, degrees o

F

 Celsius add 32 Farenheit
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