NORTH CAROLINA STATE UNIVERSITY LIBRARIES RESERVE ROOM Filename: ^lß\t^a\ •tem ID: [\^\\^Sl'^\ 'BEST COPY AVAILABLE** WARNING CONCERNING COPYRIGHT RESTRICTIONS The copyright law of the United States (Title 17, United States Code) governs the making of photocopies or other reproduction of copyrighted material. Under certain conditions specified in the law, libraries and archives are authorized to furnish a photocopy or other reproduction. One of these specified conditions is that the photocopy or reproduction is not to be used for any purpose other than private study, scholarship or research. If an electronic transmission of reserve material is used for purposes in excess of what constitutes "fair use", that user may be liable for copyright infringement. Course: NR9&F Instructor: EU?, REVIEWED The Longleaf Pine Forests of the Southeast: REQUIEM OR RENAISSANCE? VAST FORESTS OF LONGLEAF PINE (PINUS PALUSTRIS) GREETED the tust pAiropcan settlers of the lower At-lannc Coastal Plain of the Carolinas and Georgia. At that time, this species may have dominated as much as 92 million acres throughout the .southeastern United States (ľrost 1993). Only about 20 mih lion acres oř longleaf pine forest were left by 1933 (Wahlenberg 1 946). These forests declined to less than 3 million acres bv 1973, and within the next decade to 3.8 million acres (Kelly and Bechrold 1990). A more recent update (USDA borest Service, Forest Inventory and Analysis, un-publ. data) puts the total at about 3.2 million acres. Losses since then have likely reduced remaining longleaf forests to less than 3 million acres. By J. Larry Landers, . V™ oncc was one "F th5 most cxtfn" r-> -_i i i w sive forest ecosystems in North America has David H. Van Lear, and near|v vanished without notice. But only re- William D. Buyer cently has this loss begun to attract atten- Tbis naturally established second-growth longleaf pine forest, shown at age 45, arose following final harvest of the old-growth. Most second-growth loiigleaf has been cut, normally without replacement. The remnants generally range from 70 to 100 years in age. tion and concern (Means and Grow 1983; Noss 1 989). The objectives of this paper are to describe the longleaf pine ecosystem and its ecological and economic values, document its continuing decline, and explore possibilities for retaining longleaf pitre as an important part of southern forests. The Longleaf Pine Ecosystem The natural range of longleaf pine covers most of the Atlantic and Gulf Coastal Plains, from southeastern Virginia to eastern Texas and south through the northern two-thirds of Florida, with extensions into the Piedmont and mountains of north Alabama and northwest Georgia (fig. 1). The species occurs on a wide variety of sites, from wet, poorlv drained flatwoods near the coast to dry, rock\- mountain ridges (Boyer 1990). Longleaf pine is a long-lived tree, po-tentiallv reaching or exceeding an age or 300 years; however, longleaf pine foresrs Journal of Forestry 39 Table I.The distribution of longleaf pine, by state and ownership class. are often exposed to catastrophic hazards such as blowdown or fire and to continuing attrition from lightning. Most of the longleaf pine range is within 150 miles of the Atlantic or Gulf coasts, a region subject to damaging tropical storms. As a result, few longleaf pines are likely to reach their biological potential. Longleaf pine is a very intolerant pioneer species; it is a poor seed producer, and its large seeds have a limited dispersal range. Once established, seedlings grow slowly and, where vigorous competition is present, may remain in the stemless grass stage for years. The longleaf pine ecosystem is distinguished by open, park-like "pine barrens," which are composed or even-aged and multi-aged mosaics of forests, woodlands, and savannas, with a diverse groundcover dominated by bunch grasses and usually free of understory hardwoods and brush. Longleaf pine is the key tree species in a complex of fire-dependent forest ecosystems long native to the southeastern United States. Its existence was dependent on periodic fire, to which it is adapted while most of its potential competitors are nor. An exception may be very dry sites where groundcover is too sparse to prevent seeds from reaching mineral soil or competing with established seedlings. Pine barrens are known for remarkable persistence and diversity, yet they generally lack the fertile soils or layered canopies of mixed species that characterize many other diverse systems. Elements of ecosystem persistence. The ecological persistence or pine barrens is a product or long-term interactions among chmáre, fire, and the traits of key plants. Regional dom- Figitre I. Presettlement range of longleaf pine in the southeastern United States. Source: C.C Frost 1993. All National Misc. State- County/ Forest Other ownerships forest federal owned municipal ind. Farmer corp. Individual Florida 950.7 178.5 118.6 118.4 2.3 149.9 66.4 118.7 197.9 Georgia 520.2 0.0 55.3 3.5 0.0 44.7 122.8 77.5 216.4 North Carolina 255.5 12.3 57.8 37.8 0.2 20.3 43.1 21.0 63.0 Texas 45.0 11.1 0.0 0.0 0.0 22.8 0.0 5.8 5.3 South Carolina 369.0 18.7 73.4 23.2 0.0 15.4 38.1 71.6 128.6 Alabama 535.1 81.8 5.8 5.7 10.7 190.3 73.7 59.1 108.0 Mississippi 270.3 91.5 0.0 9.9 0.0 37.3 23.2 6.6 101.8 Louisiana 232.9 61.6 12.0 0.0 0.0 75.5 0.0 36.3 47.5 Total 3,178.7 455.5 322.9 198.5 13.2 556.2 367.3 396.6 868.5 Source: USDA Forest Service, Forest Inventory and Analysis data, 1994. inance by southern pines began to redevelop about 10,000 years ago with a warming trend following retreat of the last continental ice sheet. The modern longleaf pine-turkey oak (Qtiercus laevis) forests were established in north Florida about 7,800 years ago (Watts et al. 1992), and over the next 4,000 years reinvaded rhe rest of the Southeast (Delcourt and Del-court 1987). The forests of the southeastern United States, as found in historic times, were apparently in place about 5,000 years ago, with pollen records indicating 65% pine and 15% oak plus most other elements of the modern forest (Watts 1980). Undoubtedly, these forests included extensive longleaf pine stands much like those found bv the first Euro- pean travelers in the region. Thunderstorm systems with frequent lightning and rain have prevailed over this vast region, a complex of mostly sandy (quick-drying) sites that are well exposed to natural and anthropogenic disturbances. Longleaf pine and bunch grasses (e.g., wiregrass and certain bluestems) possess traits that facilitate the ignition and spread of fire during humid growing seasons (Landers 1991). Frequent fire was largely responsible for the competitive success of longleaf pine and the grasses. These keystone species exhibit pronounced fire tolerance, longevity, and nutrient-water retention that reinforce their dominance and restrict the scale of vegetation change following disturbance. Scattered longleaf pine in slash pine area transitional to south Florida communities (Sudworth 1913). Longleat pine/wiregrass and longleaf pine/bluestem mosaic of pyroclimax communities, such as savannas, sandhills, and flatwoods (Sargent 1 Frost et al. 1986 for Virginia) Longleaf pine'bluestem savanna and woodland outside the range of wiregrass. Longleaf pine/shortleaf pine/ loblolly pine/hardwoods transition area (Lockett 1870; Sargent 1884; Mohr 1897. 1901; Sudworth 1913'Harper 192^ 1928' F oM et al 19 s'^,si/ 40 November 1995 Frequent fires during the growing season largely prevent species native to other habitats from encroaching into pine barrens. The chronic fire regime also maintains soil structure and nutrient dynamics to which longleaf pine is adapted, while inhibiting the processes of soil weathering and building (McKee 1982). These fire effects tend to make longleaf pine sites more favorable to resident species than those indigenous to more nutrient-rich habitats. Elements of ecosystem diversity. Combinations of disturbance and site factors play important roles in maintaining biological diversity. Variability is added to the disturbance regime by frequent lightning strikes, treefalls, and various animal influences that have local effects. More vast areas can be severely affected every few decades by tropical storms and hydrologie extremes. Disturbances acting across site gradients provide temporary habitat features (e.g., dead wood, hardwood thickets, bare mesic/wet patches) along with more stable features (e.g., living old pines, treeless spaces, swards, bare xeric patches). Thus, many organisms are adapted to this disturbance-prone but relatively stable environment. Among them are opportunists coexisting in habitat mosaics with longer lived species. Many birds and mammals use cavities made by the red-cockaded woodpecker, and at least 60 vertebrate and 302 invertebrate species use gopher tortoise burrows (Jackson 1989). Thus, spatial and temporal variations in habitat, coupled with commensal relationships, accommodate an unusual variety of species for a given area. The diversiry of groundcover plants per unit area places longleaf pine ecosystems among the most species-rich plant communities outside the tropics (Peet and Allard 1993). Yet longleaf pine ecosystems now occupy only a small part of their original area. Extreme habitat reduction is the main cause for the precarious state of at least 191 taxa of vascular plants (Hardin and White 1989; Walker 1993) and such key wildlife as the red-cockaded woodpecker, gopher tortoise, and southern fox squirrel. A concerted and committed effort to restore and manage longleaf pine ecosystems would help ensure a future for this important part of the nation's natural heritage. Decline of the Longleaf Forest Because of its many desirable attributes, longleaf pine has been intensively exploited ever since colonial times (Croker 1979). In early settlement days, trees were cut to build cabins and homes or to clear land for crops. Because waterways formed the major routes for transportation through most of the 19th century, timber, usually larger trees, was cut where it could be easily moved to water. The advent of railroad logging near the end of the 19th century provided access to the remaining longleaf timberland. Merchantable trees were cut, and much of the remainder damaged or destroyed. Cutting generally proceeded from the Atlantic states west through the Gulf Coast region to Louisiana and Texas, with increasing intensity of use as time went on. Long-leaf pine logging reached a peak in 1907, when an estimated 13 billion board feet were cut (Wahlenberg 1946). Logging came to a halt with the Great Depression and the near exhaustion of the timber resource. Second-growth longleaf pine, to varying degrees, succeeded old-growth on millions of acres throughout the Southeast. Cutting in this natural second-growrh forest began during World War II and continues today as this age class has matured. Over 90% of the longleaf pine remaining in 1985 was natural in origin (Kelly and Bechtold 1990). Annual removals exceeded growth by 43%. Declines were observed in every diameter class below 16 inches while increases were confined to the 16-inch and larger diameter classes, suggesting that most remaining foresrs are aging without replacement. A combination of circumstances appears responsible for the long-term decline of the longleaf pine forest. Much of the land in the region was once cleared for cropland or pasture. Longleaf pine does not successfully invade open land in competition with more aggressive species. It has sometimes succeeded old-growth when periodic fires provided a seedbed and controlled woody competition, and when wild hogs did not reach a density high enough to destroy established seedlings. The disruption of natural fire regimes, resulting in part from forest fire protection policies implemented during the 1920s, however, allowed invasion of longleaf sites by hardwoods and more aggressive southern pines. Longleaf pine and its associated community cannot naturally succeed itself under these conditions. Since the end of World War II, many large lumber companies that owned and generally favored longleaf pine for high quality logs and poles have slowly been ac- Figure 2. Current distribution of longleaf pine, by county, in the southeastern United States. Source: USDA Forest Service 1994. Map produced by Donald van Blaricom, Clemson University. Journal of Forestry 41 THE WORLD AWAITS YOU. As a forestry professional, you've got knowledge and practical experience that are in demand in developing countries all over the world. As a Peace Corps Volunteer, you can share your skills and make a lasting difference in people's lives. Peace Corps offers two years of international experience, language and cross-cultural training, paid living expenses, medical care, opportunities for graduate school and more. It's an unforgettable opportunity for personal and career growth! Peace Corps Volunteers must be U.S. citizens. There is no upper age limit. Married applicants are welcome, but your spouse must also qualify for an assignment. TO LEARN MORE, PLEASE CALL: 800-424-8580 MAIL TO: Name. Peace Corps, Room 8506 1990 K Street, NW Washington, DC 20526 Address. City------- Phone# ( )___________________ U.S. citizen: □ Yes □ No Please describe your forestry experience: State — SSN#_ Age_ ZIP- Privacy Act Notice: Providing the information herein required is voluntary. It will only be used to determine your interest in Peace Corps programs. Auth: 22USC 2501 0MB Approval number 0420-0007. Estimated completion time: 1.75 minutes. Send comments on this estimate to Office of Management and Budget, Project 0420-0007, Washington, DC 20503 or to Peace Corps, Paperwork Reduction Project 0420-0007, Washington, DC 20526. Exp. 1/31/97. 724 quired by the pulp and paper industry. These companies' need to maximize volume yield over the shortest possible period of time led to intensive short-rotation plantation management using genetically improved loblolly (Pinus taeda L.) or slash pine (P. elliotti). Longleaf pine is often considered a slow-growing species that is difficult to regenerate and that cannot economically compete with the other pines. Loblolly and slash pine are thus the preferred species for planting and recently accounted for 95% of all southern pine seedling production. Management Options Longleaf pine has many attributes that allow a variety of management options. It has always been recognized as a high-quality timber tree providing a wide range of products: logs, poles, piling, posts, peelers for plywood, and pulpwood. It almost always has a higher specific gravity than the other southern pines and thus produces more dry weight per unit of volume. On average sites, 30 to 80% of the trees in a longleaf stand will make poles, which are more valuable than sawlogs (Boyer and White 1990). In addition to its commercial quality and versatility, longleaf pine, once established, is a low-risk species to manage. It is resistant to fire and the more serious diseases and insect pests that afflict other southern pines, including fusiform rust, annosus root rot, pitch canker, southern pine beetle, and tipmoth. Longleaf pine is more resistant than slash pine to breakage from ice storms (Van Lear and Saucier 1973). The species develops a massive taproot that, in mature trees, may reach a depth of 8 to 12 feet or more, helping reduce the risk of windthrow (Boyer 1990). It is thus suited to a wide range of management goals and silvicultural methods. These include even-aged, two-aged, and, on many sites, a range of all-aged management methods (Farrar and Boyer 1991). Because of longleaf pine's natural tolerance of fire, uneven-aged stands can be regularly burned at 2- to 4-year intervals to control hardwoods and brush or prepare a seedbed without the need for any special measures to protect regeneration. The rapid early breakup of longleaf seedling stands into a range of size classes helps reduce the risk of excessive seedling mortality from untimely fire. In addition, longleaf pine's reputation as the slowest growing southern pine may be largely undeserved. On many former long-leaf sites, based on side-by-side comparisons, the species grows as well as, .or better than, the other southern pines once it has emerged from the grass stage (Schmidtling 1987; Outcalt 1993). If desired, longleaf pine on average sites can produce poles and logs in 40- to 50-year rotations, possibly less in plantations. Most growth data for natural longleaf pine have been derived from forests that periodically burn. Both the diameter and height growth of young longleaf pines are reduced by regular burning (Boyer 1994). The reduction in height growth alone is equivalent to a loss of 5 feet in site index. The growth reduction associated with periodic fires has been regularly reported over the years, and the bias this introduces for site classification and yield table construction has long been recognized (Cary 1932). Many landowners, however, may be willing to accept lower yields in return for the natural beauty and enhanced biodiversity of open, regularly burned, longleaf forests. Longleaf pine can be naturally or artificially regenerated at a reasonable cost and with a high probability of success if necessary cultural measures are properly timed and executed. Quality planting stock along with appropriate cultural treatments can reduce the grass stage to one or two years. Longleaf pine straw is also becoming a major and valuable product in many areas. Pine-straw raking can be detrimental to legumes and other species in the herb layer, however, and should be avoided where species diversity is valued (Frost 1990). Permanently reserving some areas from raking may preserve species diversity by providing local refugia. Pine-straw production peaks at about the time that volume increment culminates, so a regulated forest will allow pine straw to be efficiently gathered on a continual basis, even if harvesting is confined to a single age class for a relatively short period of time. Fertiliza-, tion can make up for the nutrients removed with the straw. Many game species, ranging from white-tailed deer and wild turkey to bob-white quail and rabbit, thrive in longleaf pine forests, especially those maintained in an open condition by frequent thinnings and prescribed fire. These open habitats are in high demand by hunters and provide landowners the opportunity to lease their lands to sports enthusiasts. Fee hunting provides a means of allocating access to public resources on private lands. Another option in longleaf pine management is woodland grazing for beef cattle. Longleaf pine forests kept open (basal 42 November 1995 9 CRAWFORD AND BOURLAND Forestry, Wildlife and Environmental Consultants • Forest Management Plans • Forest Product Sales • Timber Estimates * Appraisals • Integrated Wildlife Management • Hunting Lease Management • Wildlife Habitat Development • Site Analysis • Endangered Species • Wetlands Sam D. Crawford, A.C.F. Tom Bourland Consulting Forester Certified Wildlife Biologist Crawford and Bourland, Inc. 2529 East 70th, P.O. Box 5933, Shreveport, LA 71135-5933 Ph. (318) 797-1546 FAX (318) 797-1545 areas 50 to 70 ft2/ac or less) by prescribed fire and thinning maintain relatively high forage yields throughout the timber rotation. For longleaf pine-wiregrass ranges, late winter or early spring burns on a two-year rotation work well after pines are large enough to tolerate fire. Restoration of the Longleaf Pine Ecosystem One of the keys to restoration of the longleaf pine ecosystem is to ensure that its recovery benefits society in general and private landowners in particular. Without economic benefits'long-term and broad-scale conservation projects are usually doomed to failure (Kimmins 1992; Oliver 1992). Harvesting need not be eliminated or even moderately restricted to restore and maintain longleaf pine ecosystems, as evidenced by the fact that logging at the turn of the century apparently had little effect on groundcover diversity (Noss 1989). Any restrictions on harvest would be a disincentive to many landowners and could result in elimination of much of the remaining longleaf on private lands. Restoration appears achievable. As seen from its present distribution (fig. 2), long-leaf pine still occurs over most of its former range, albeit in remnant pockets. By expanding these pockets, it should be feasible to gradually expand longleaf pine acreage. Education, research, commitment on the part of resource managers, and a landscape perspective are essential ingredients in any such effort. Education of the public regarding the current status of the longleaf pine ecosystem, its potential economic value, its outstanding biodiversity, and the role of fire in maintaining the system is an initial step in securing support for restoration. With public support, federal, state, and local laws can be changed to provide restoration incentives. Foresters and other resource managers, in partnership with conservation groups, can also promote the use of fire as an ecological force necessary for maintenance of fire-dependent natural communities. Frequent prescribed burning, including the use of growing-season fires where appropriate, promotes the diversity and stability of these communities (Noss 1989; Frost 1990). In 1990, Florida passed a Prescribed Burning Act authorizing and promoting prescribed burning for ecological and other purposes. As other southern states follow suit, the ability to use this tool for restoring and maintaining longleaf pine ecosystems will be greatly enhanced. Federal and state agencies, universities, forest industries, and research organizations are beginning to cooperate and commit to research and technology transfer promoting the ecological and economic attributes of the longleaf pine ecosystem. Technology is now available that allows regeneration of longleaf pine to approach parity with the other southern pines. Mechanical site preparation techniques, however, should be carefully evaluated if adverse effects on associated species such as wiregrass are to be avoided (Clewell 1989). Natural regeneration methods using a regime of frequent fire are already compatible with maintenance of these communities (Boyer and White 1990). While many private landowners are concerned about the environment and the amenities their land provides, most are also motivated by the need to generate income from their land. Longleaf pine can be managed in an ecologically sensitive manner that generates a satisfactory income (Landers et al. 1990). Agencies administering public cost-share programs, such as the Forest Incentives Program and the Stewardship Incentives Program, could promote longleaf pine regeneration. Many of the actions suggested by Oliver (1992) to resolve pressures facing Pacific Northwest forests would also be useful in restoring the long-leaf pine ecosystem. For example, subsidies or tax deferrals could provide longleaf pine landowners with incentives to regenerate longleaf pine and maintain the balance of age classes and stand structures necessary to protect plant and animal diversity across the landscape. Longleaf pine occurs on. approximately 3.2 million acres (table 1). Almost two-thirds of this acreage is in Florida, Alabama, and Georgia. North Carolina, South Carolina, Mississippi, and Louisiana each contain more than 200,000 acres. About 31% of the longleaf pine acreage is owned by public agencies, 18% by forest industry, and the remaining 51% by a variety of private owners, most of them not farmers. This mix of ownerships emphasizes that effective restoration will require a multi-owner approach across the landscape. A balanced longleaf pine forest, supplementing other forest types, could help achieve greater productivity and stability throughout the forest economy of the Southeast. Restoration of small, fragmented stands of longleaf pine is helpful but will not suffice to sustain a functioning ecosystem. The challenge is to restore this eco- Journal of Forestry 43 SOFTWARE FOR ENDANGERED ^ SPECIES JM Since 1982 JmlS RAMAS® $j^^ Our models and software are in use in over 400 institutions in 20 countries. Our recent projects have considered: • Spotted owl in Washington, • Gnatcatcher in California, • Red-cockaded woodpecker in Louisiana. CALL FOR A FREE CATALOG tel (800) 735-4350 fax (516) 751-3435 Applied Biomathematics 100 North Country Road Setauket,NY 11733 USA system in an economically viable way so that private landowners can be voluntary partners who benefit in the process. Existing disincentives and longstanding biases must be overcome and incentives added, so that landowners are encouraged to regenerate and manage longleaf pine using methods that simulate historical processes. Restoration of longleaf pine forests should not be viewed as competition for intensive pine plantation management so common in the South. Intensive forestry aimed at high productivity is also an essential component of the southern landscape. By maintaining high yields, intensive plantation management reduces pressure on other parts of the landscape, allowing other land uses such as longleaf pine ecosystem restoration. In turn, diverse and productive longleaf pine forests, growing large, high-value products under relatively long rotations, would complement the economic contribution made by other species while preserving environmental values that are a part of the heritage of the southern landscape. The future will likely reveal unforeseen values in native forest ecosystems that will greatly benefit society and translate into additional economic incentives for landowners. Restoring the longleaf pine ecosystem could serve as a prime example of forest ecosystem management—how a once diminished ecosystem was restored as a sustainable, functioning paradigm through wise stewardship. E33 Literature Cited BOYER, W.D. 1990. Pinus palustris Mill, longleaf pine. In Silvios of North America. Vol. 1, Conifers, tech. coords. R.M. Burns and B.H. Hon-kala.Washington, DC: USDA Forest Service. ---------. 1994. Eighteen years of seasonal burning in longleaf pine: Effects on overstory growth. In Proceedings of the 12th Conference on Fire and Forest Meteorology. Bethesda, MD: Society of American Foresters. BOYER, W.D., and J.B. White. 1990. Natural regeneration of longleaf pine. In Proceedings of the Symposium on the Management of Longleaf Pine, ed. R.M. Farrar Jr. General Technical Report SO-75. New Orleans: USDA Forest Service, Southeastern Forest Experiment Station. Cary, A. 1932. Some relations of fire to longleaf pine. Journal of Forestry 30:594-601. Clewell, A.F. 1989. Natural history of wiregrass (Aristida stricta Michx., Gramineae). Natural Areas Journal9:223-33. Croker, T.C., Jr. 1979. Longleaf pine: The long-leaf pine story. Journal of Forest History 23:32-43. Delcourt, P.A., and H.R. Delcourt. 1987. Long-term forest dynamics of the temperate zone. New York: Springer-Verlag. Farrar, R.M.,Jr„ and W.D. Boyer. 1991. Managing longleaf pine under the selection system — promises and problems. In Proceedings, Sixth Biennial Southern SilviculturalResearch Conference, eds. S.S. Coleman and D.G. Neary. General Technical Report SO-75. New Orleans: USDA Forest Service, Southeastern Forest Experiment Station. FROST, C.C. 1990. Natural diversity and status of longleaf pine communities. In Forestry in the 1990s—A changing environment, eds. G. Youngblood and D.L. Frederick. Bethesda, MD: Society of American Foresters. ---------. 1993. Four centuries of changing landscape patterns in the longleaf pine ecosystem. Proceedings, Tall Timbers Fire Ecology Conference 18. Tallahassee, FL: Tall Timbers. Hardin, E.D., and D.L. White. 1989. Rare vascular plant taxa associated with wiregrass (Aris-^ tida stricta) in the southeastern United States. Natural Areas Journal9:234—45. JACKSON, D.R. 1989. The fauna of gopher tortoise burrows. In Proceedings of the Gopher Tortoise Relocation Symposium, eds. J.E. Diemer, et al. Technical Report 5. Tallahassee: Florida Game and Fresh Water Fish Commission, Nongame Wildlife Program. Kelly, J.F., andWA. Bechtold. 1990. The long-leaf pine resource. In Proceedings of the Symposium on the Management of Longleaf Pine, ed. R.M. Farrar Jr. General Technical Report SO-75. New Orleans: USDA Forest Service, Southeastern Forest Experiment Station. KlMMINS, J.P. 1992. Balancing act: Environmental issues in forestry. Vancouver, BC: UPC Press. LANDERS, J.L. 1991. Disturbance influences on pine traits in the southeastern United States. Proceedings, Tall Timbers Fire Ecology Conference 17. Tallahassee, FL: Tall Timbers. Landers, J.L., N.A. Byrd, and R. Komárek. 1990. A holistic approach to managing longleaf pine communities. In Proceedings of the Symposium on the Management of Longleaf Pine, ed. R.M. Farrar Jr. General Technical Report SO-75. New Orleans: USDA Forest Service, Southeastern Forest Experiment Station. McKee, W.H. 1982. Changes in soil fertility following prescribed burning on Coastal Plain pine sites. Research Paper SE-234. Asheville, NC: USDA Forest Service, Southeastern Forest Experiment Station. MEANS, D.B., and G. Grow. 1985. The endangered longleaf pine community. ENFO 85(4): 1-12. NOSS, R.F. 1989. Longleaf pine and wiregrass: Keystone components of an endangered ecosystem. Natural Areas Journal 9:234-35. OLIVER, CD. 1992. Achieving and maintaining biodiversity and economic productivity. Journal of Forestry 90(9):20-25. OUTCALT, K.W. 1993. Southern pines performance on sandhill sites in Georgia and South Carolina. Southern Journal of Applied Forestry 17:100-02. Peet, R.K., and D.J. Allard. 1993. Longleaf pine-dominated vegetation of the southern Atlantic and eastern Gulf Coast region, USA. Proceedings, Tall Timbers Fire Ecology Conference 18. Tallahassee, FL: Tall Timbers. SCHMIDTLING, R.C. 1987. Relative performance of longleaf compared to loblolly and slash pines under different levels of intensive culture. In Proceedings of the Fourth Biennial Southern Silvicultural Research Conference, comp. D.R. Phillips. General Technical Report SE-42. Asheville, NC: USDA~Forest Service, Southeastern Forest Experiment Station. Van Lear, D.H., andJ.R. Saucier. 1973. Comparative glaze damage in adjacent stands of slash and longleaf pine. Forest Research Series, 27. Clemson, SC: Clemson University. WAHLENBERG, W.G. 1946. Longleaf pine: Its use, ecology, regeneration, protection, growth, and management. Washington, DC: C.L. Pack Forestry Foundation and USDA Forest Service. WALKER, J.L. 1993. Rare vascular plant taxa associated with the longleaf pine ecosystem. Proceedings, Tall Timbers Fire Ecology Conference 18. Tallahassee, FL: Tall Timbers. Watts, W.A. 1980. The late quaternary vegetation history of the southeastern United States. Annual Review of Ecology and Systematics 11:387—409. Watts, W.A., B.C.S. Hansen, and E.C. Grimm. 1992. Camel Lake: A 40,000-year record of vegetational and forest history from north Florida. Ecology 73:1056-66. ABOUT THE AUTHORS The late J. Larry Landers was conservation coordinator, Joseph W. Jones Ecological Research Center at Ichauway, Newton, GA; David H. Van Lear is Bowen Professor of Forestry, Department of Forest Resources, Clemson University, SC'29634-1003; William D. Boyer is research forester, Southern Research Station, USDA Forest Service, Auburn University, AL. 44 November 1995